Messinian salinity crisis record under strong freshwater input in marginal, intermediate, and deep environments: The case of the North Aegean

Accepted Manuscript Messinian salinity crisis record under strong freshwater input in marginal, intermediate, and deep environments: The case of the North Aegean

Vasileios Karakitsios, Jean-Jacques Cornée, Theodora Tsourou, Pierre Moissette, George Kontakiotis, Konstantina Agiadi, Emmanouil Manoutsoglou, Maria Triantaphyllou, Efterpi Koskeridou, Harikleia Drinia, Dimitrios Roussos PII: DOI: Reference:

S0031-0182(17)30116-5 doi: 10.1016/j.palaeo.2017.06.023 PALAEO 8339

To appear in:

Palaeogeography, Palaeoclimatology, Palaeoecology

Received date: Revised date: Accepted date:

2 February 2017 7 June 2017 16 June 2017

Please cite this article as: Vasileios Karakitsios, Jean-Jacques Cornée, Theodora Tsourou, Pierre Moissette, George Kontakiotis, Konstantina Agiadi, Emmanouil Manoutsoglou, Maria Triantaphyllou, Efterpi Koskeridou, Harikleia Drinia, Dimitrios Roussos , Messinian salinity crisis record under strong freshwater input in marginal, intermediate, and deep environments: The case of the North Aegean, Palaeogeography, Palaeoclimatology, Palaeoecology (2017), doi: 10.1016/j.palaeo.2017.06.023

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ACCEPTED MANUSCRIPT Messinian salinity crisis record under strong freshwater input in marginal, intermediate, and deep environments: the case of the North Aegean KARAKITSIOS Vasileios1, CORNÉE Jean-Jacques2, TSOUROU Theodora1, MOISSETTE Pierre3,1, KONTAKIOTIS George1, AGIADI Konstantina1, MANOUTSOGLOU Emmanouil4, TRIANTAPHYLLOU Maria1, KOSKERIDOU Efterpi1, DRINIA Harikleia1,

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ROUSSOS Dimitrios1

National and Kapodistrian University of Athens, Faculty of Geology and Geoenvironment,

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Department of Historical Geology & Paleontology, 15784, Athens, Greece,

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[email protected]

Géosciences Montpellier, CNRS, Université de Montpellier, Montpellier, France

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Univ Lyon, Université Lyon 1, Ens de Lyon, CNRS, UMR 5276 LGL-TPE, 69622

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Villeurbanne, France 4

Technical University of Crete, School of Mineral Resources Engineering, GR - 731 00

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Chania, Greece

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Abstract

In the present study, we investigate the Mediterranean–Paratethys connection during

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the late Miocene in Strymon Basin (North Aegean, northeastern Mediterranean) and compare this onshore sequence with the adjacent offshore Prinos-Nestos sequence, before, during, and after the Messinian Salinity Crisis (MSC). Strymon Basin was a peripheral shallow-water

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basin during the first MSC stage. The Akropotamos sections expose a clastic sequence with gypsum intercalations, which is dated in the Messinian based on the ostracod and calcareous

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nannofossil assemblages. This sequence records the Primary Lower Gypsum deposition in a shallow marine environment and its passage via the Messinian erosional surface to a brackish environment with changing salinity conditions similar to the Paratethyan depositional environments. The sequence is capped by a travertine marker horizon observed across the entire Strymon Basin indicating freshwater environment. The Miocene–Pliocene transition is characterized by salinity changes caused by the interaction between Atlantic-Mediterranean and Paratethyan waters, predating the marine reflooding at the end of the MSC, which is attested by the overlying Pliocene open marine deposits. The offshore Prinos-Nestos basin incorporates the Nestos intermediate basin and the Prinos intermediate-deep basin. Borehole and seismic profile data from the offshore Prinos-Nestos oil field reflect a thick clastic 1

ACCEPTED MANUSCRIPT sequence, topped by turbidites, and followed by an evaporitic unit deposited during the Messinian. In the Nestos slope area, the evaporite unit consists of anhydrite-shale alternations. Toward the basin’s depocenter (Prinos Basin), anhydrite is replaced by halite. The sequence is overlain by Pliocene–Holocene deltaic prograding deposits. Sedimentologic and biostratigraphic data show that the thick halite-shale couplets in the deepest part of the offshore Prinos-Nestos Basin were deposited under permanent marine conditions, suggesting no desiccation before, during, and after the MSC. Conclusively, the present results indicate

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that a connection between the Mediterranean and the Paratethys was occasionally established before the Pliocene reflooding and favor the non-desiccation MSC model for the deep marine

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evaporite deposition.

Keywords: Akropotamos; travertine; oil and gas field; North Aegean Sea; evaporites;

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desiccation

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1. Introduction

The late Miocene connection between the Mediterranean Sea and the Paratethys

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brackish basin (Fig. 1) has been investigated since the last century (Hsü et al., 1977, 1978; Hsü, 1972, 1978; Rögl and Steininger, 1983). A hypothesis was initially formulated that a

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gateway or a drainage network was installed through the Balkans before and after the Messinian salinity crisis (MSC). Further research revealed marine calcareous nannoplankton in the Pontian deposits of the Dacic Basin (Mǎrunţeanu and Papaianopaol, 1995; Snel et al.,

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2006a) and in the late Messinian deposits of the Strymon Basin, northern Greece (Snel et al., 2006b). In addition, periodic exchanges were suggested between the Mediterranean and the

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Dacic Basin during late Messinian sea-level highstands (Clauzon et al., 2005; Popov et al., 2006; Popescu et al., 2009; Suc et al., 2011; Bache et al., 2012; Krstić et al., 2012; Bakrač et al., 2012; Do Couto et al., 2014; Suc et al., 2015; Kontakiotis et al., 2016; Agiadi et al., 2017). Overall, four areas were identified as possible Mediterranean–Paratethys gateways: the Bosporus straits (Popov et al., 2006), the Strymon Basin (Snel et al., 2006b; Suc et al., 2015; Figs. 1 and 2), SE Turkey connecting the eastern Mediterranean to the Black Sea (van Baak et al., 2016), and a connection between the eastern Mediterranean and the Caspian Sea passing through northern Iran (van Baak et al., 2016). Calcareous nannoplankton and planktonic foraminifers showed that, during high sealevel episodes prior and after the MSC, the Mediterranean and Dacic Basin were connected 2

ACCEPTED MANUSCRIPT (Clauzon et al., 2005; Snel et al., 2006a,b; Suc et al., 2011, 2015). The crossed exchanges of organisms between the Mediterranean and Dacic Basin through the Balkans marine gateway during high sea-level episodes were tested to explain the Lago Mare deposits (Clauzon et al., 2005; Suc et al., 2011; Bache et al., 2012). According to Snel et al. (2006a,b), the nannofossil assemblages show that temporary connections of Atlantic/Mediterranean waters with the Northern Aegean Basin existed in the latest Messinian Strymon Basin, and these marine ingressions likely extended further into the Eastern Paratethys in the middle and upper

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Pontian of the Dacic and Euxinic Basins. The same authors point out that they did not find indications for a tectonically induced transgression caused by ongoing subsidence of the

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Pliocene; the presence of brackish-water faunas and intercalated nannofossil assemblages in

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the uppermost part of the Messinian illustrates that, before the lower Pliocene, the basin was not completely dessicated. Nevertheless, the Miocene–Pliocene transition in this area was

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characterized by a salinity change, reflecting the increased influence of Atlantic water. Suc et al. (2015) attributed the Mediderranean-Paratethys marine corridor in the Strymon and intraBalkans area to the regional tectonic extension, whereas the tectonically controlled subsidence

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caused the corridor’s closure creating small lagoons and lakes around the corridor during the high sea-level phases. According to these authors, during the peak of the MSC, the corridor

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was probably evolved as a powerful river, contributing to the deposition of clastics in the Prinos-Nestos hydrocarbon field. This interpretation suggests a significant erosional

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unconformity (MES) during the peak of the MSC. The above interpretations show the controversial opinions concerning the record of the MSC and the related processes. However, it should also be taken into account that the onshore deposits cannot fully

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reveal the MSC extent and processes. Indeed, MSC onshore deposits are often difficult to correlate with offshore sequences, either because they are not contemporary or because they

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are tectonically disconnected from them. Most of the onshore MSC evaporites were accumulated in shallow-water marginal/peripheral basins. These deposits predate the drawdown phase and are not coeval to the Messinian successions of the deep Mediterranean basins, which probably accumulated while the marginal/peripheral basins deposits were eroded. Consequently, the marginal/peripheral deposits are incomplete, because they underwent subaerial or submarine erosion during the drawdown phase (Lofi et al., 2011a,b; Roveri et al., 2014a). On the contrary, the deep offshore MSC basins, where the largest volume of MSC evaporites and clastic products were accumulated, were possibly never fully desiccated and present complete successions recording the drawdown phase. MSC scenarios are rarely based on observations from the offshore domain, mainly because there are no 3

ACCEPTED MANUSCRIPT boreholes penetrating the entire evaporitic unit (Roveri et al., 2014a); hence, the offshore MSC record is only inferred from seismic profiles and rare borehole data covering only the upper part of the event sequence. Therefore, the resolution of lateral correlations between onshore/offshore data is limited. In addition, the evaporative concentration model referring to the total-desiccation hypothesis of the Mediterranean deep basins during the MSC is not adequately supported by facts (Roveri et al., 2016). The studied area in northern Greece (Figs. 1, 2, and 3) is crucial, because the onshore

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Strymon Basin probably constituted a corridor between the Mediterranean and the Paratethys, whereas the adjacent offshore Prinos-Nestos Basin contains a thick evaporitic unit, which has

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been drilled throughout many times across the basin for hydrocarbon exploration and

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exploitation. In the present study, we provide new biostratigraphic (ostracods and calcareous nannofossils), paleoecologic (benthic foraminifers, bivalves, and ostracods), sedimentologic,

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and field data, which elucidate aspects of the Balkan marine gateway between the Mediterranean and the Dacic Basin. Furthermore, we exploit the available offshore data on the Messinian evaporites across the Prinos-Nestos Basin, correlating it with the onshore

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Strymon Basin.

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2. Geological setting

The North Aegean region consists of the alpine Vardar zone, and the Serbo-

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Macedonian and Rhodope massifs (inset map of Fig. 2). The tectonic evolution of this region was explained in the context of the North Aegean core complexes (Fig. 2), which were developed soon after the end of the continental block convergence and the piling up of the

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Hellenic thrust wedge (Dinter, 1988; Dinter and Royden, 1993; Kydonakis et al., 2015). The core complexes were formed due to back-arc extension, driven by the Hellenic slab rollback

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behind the thrust wedge. The middle Miocene–lower Pliocene deposits, which filled the basins due to the extension in the back-arc of the Hellenic subduction zone, were overlaid by late Pliocene–Quaternary deposits. The latter filled the North Aegean basins that were created during the late Pliocene in response to strike-slip movements in the North Aegean trough, due to the movements of the North Anatolian Fault (Dinter, 1988; Roussos and Lyssimachou, 1991; Dinter and Royden, 1993; Kydonakis et al., 2015). The Strymon Neogene Basin (northern Greece), at the southern edge of the Rhodope Massif, is one of the extensional basins that were developed in the early–middle Miocene over the southern part of the northern Aegean along NE-SW and NW-SE trending faults (Figs. 2 and 3) associated with the above mentioned tectonic evolution (Dinter, 1988; Dinter 4

ACCEPTED MANUSCRIPT and Royden, 1993; Brun and Sokoutis, 2007; Wüthrich, 2009; Burg, 2012; Kydonakis et al., 2015). Akropotamos area is located in the southeastern part of the Strymon Basin and comprises (Figs. 2 and 3) more than 700 m of Neogene deposits. The lower part of the Neogene sequence (more than 300 m) is composed of non-marine fluvio-lacustrine clastic deposits and rare clays with lignite intercalations. These are followed by 80 m well-bedded clastic and carbonate alternations corresponding to lacustrine, brackish, or shallow marine

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environments. Above lies a relatively open marine sandy or silty clay unit (approximately 40 m thick), locally intercalated by lens-shaped gypsum bodies. Subsequently,

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lacustrine/brackish beds are topped by travertinous limestones, which are followed by lower

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Pliocene open marine clastics and marl alternations (Dermitzakis et al., 1986; Karistineos and Georgiades-Dikeoulia, 1986). This sequence is unconformably covered by upper Pliocene–

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Pleistocene shallow marine sands and gravels (Steffens et al., 1979; Psilovicos and Syrides, 1983; Dinter and Royden, 1993).

The Neogene deposits of the offshore Prinos-Nestos Basin (Figs. 2 and 3) comprise a

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clastic succession of 4000 to 5000 m thick (Lalechos and Savoyat, 1977; Pollak, 1979; Proedrou, 1979, 1988; Proedrou and Sidiropoulos, 1992; Proedrou and Papaconstantinou,

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2004). The oldest Neogene sediments were penetrated by the Peramos well (Fig. 3B) and are of Langhian–Serravallian age. In the Prinos-Nestos Basin, the maximum thickness of the pre-

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evaporitic unit is approximately 2100 m (Proedrou and Sidiropoulos, 1992). The sequence within the Prinos deep area of the basin (Fig. 4) commences with continental deposits, followed by thick marine shale deposits (constituting the oil source rocks of the Prinos field)

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with carbonate and sandstone intercalations. In this succession, salt and anhydrite deposits are observed in three stratigraphic intervals (Proedrou and Sidiropoulos, 1992). The first interval,

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40–50 m thick, is located close to the basement. The second is about 40 m thick and includes some dolomite. It is found just before the 300-m thick Prinos turbidite deposits, which constitute the main hydrocarbon reservoirs of the Prinos oil field. After 15 m of marine shales on top of the turbidites follows the third, main evaporitic sequence, constituting the Messinian evaporitic unit, with a thickness ranging from 600 to 800 m. Cores from the Prinos area showed that it is composed of seven halite-shale couplets in which the lowest salt layer is the thickest (60–100-m thick) and best developed. Anhydrite clastics are present at the base of the lower salt layer. These clastics are gradually increasing toward the Nestos slope area (Proedrou, 1979). Cores from the Nestos slope area of the basin showed that the evaporitic unit is composed by anhydrite layers alternating with well-bedded shales, claystones, 5

ACCEPTED MANUSCRIPT turbidite/conglomerate, and rare carbonate layers, and the total thikness of the unit is between 500 and 600 m, whereas the equivalent to the Prinos turbidite deposits corresponds to a prodeltaic zone (Lalechos and Savoyat, 1977; Pollak, 1979; Proedrou, 1979; Proedrou and Papaconstantinou, 2004). The post-evaporitic sequence of the Prinos-Nestos Basin was deposited in the Pliocene–Pleistocene (Pollak, 1979; Proedrou, 1979; Sidiropoulos, 1980). It consists mainly of poorly sorted sands and clays reaching a maximum thickness of about 2700 m in the

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present center of the basin. The lower Pliocene deposits of the basin’s center are open marine shales, passing toward the basin slope to shallow marine deposits, indicating that marine

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sedimentation was taking place concurrently over the entire basin. The top coarse clastic

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deposits with abundant mollusks point to a deltaic prograding sequence, followed again by transgressive marine clastic sediments (Proedrou and Papaconstantinou, 2004; Kiomourtzi et

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al., 2007).

Biostratigraphic data from Sidiropoulos (1980) for the slope area revealed in the Nestos-2 well the presence of the calcareous nannofossil Discoaster quinqueramus in the

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shale intercalations 40 m below the top of the evaporitic unit and the continuous presence of the benthic foraminifer Uvigerina rutila in the stratigraphic level 60 m above the evaporitic

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unit (Fig. 4). In Nestos-1 and 2 wells, Sidiropoulos (1980) also recorded the occurrence (for about 15 m) of the planktonic foraminifer Globorotalia puncticulata in the stratigraphic level

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located 330 m above the base of the post-evaporitic unit and the presence of the planktonic foraminifer Turborotalita quinqueloba (clarkei) and the nannofossils Coccolithus pelagicus

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and Braarudosphaera bigelowii in the shale alternations of the evaporitic unit (Fig. 4).

3. Materials and methods

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The Neogene infill of the Strymon Basin (Fig. 2) was studied south of Akropotamos village, about 45 km WSW of the Kavala City (Figs. 2 and 3). The study of the Neogene sediments’ distribution was based on tectonic, stratigraphic, paleontologic, and sedimentologic observations. Field observations were combined with detailed sampling in four sections revealed in three abandoned gypsum quarries. The biostratigraphic and paleoecologic data were obtained by analyzing ostracods, benthic foraminifers, invertebrates (mollusks), and calcareous nannofossils, as planktonic foraminifers were not found. In particular, 61 samples were obtained for the paleontological analysis. Samples with the prefix AK came from Section A, those with BK from Section B, those with BK’ from Section C, and those with CK from Section D (Fig. 5). The samples 6

ACCEPTED MANUSCRIPT were disaggregated with a 5% H2O2 solution, washed over 0.125 mm and 0.063 mm mesh sieves, and the residues were oven-dried at approximately 40°C. Furthermore, we exploited the data of the published works (boreholes and seismic profiles) from the Prinos-Nestos Basin, focusing on oil research, in terms of their significance for the Messinian salinity crisis. These data were obtained from offshore exploratory wells that were drilled from 1976 to 2015 in order to investigate the Prinos-Nestos hydrocarbon field and the future petroleum and gas perspectives of the adjacent areas (Lalechos and

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Savoyat, 1977; Pollak, 1979; Proedrou, 1979, 1988; Georgakopoulos et al., 1991; Proedrou and Sidiropoulos, 1992; Georgakopoulos, 1998; Proedrou and Papaconstantinou, 2004;

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Kiomourtzi et al., 2007; 2008).

3.1 Ostracods

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Only 19 out of the 61 samples revealed ostracod assemblages; most of the samples were barren. The ostracods were handpicked from the residue fraction greater than 0.125 mm, and they were identified using a Leica stereomicroscope and a scanning electron microscope

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(Jeol JSM 6360; University of Athens, Faculty of Geology and Geoenvironment). The ostracod remains mainly included carapaces; loose valves were rare. A semi-quantitative

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analysis was performed on the ostracod faunas; each carapace was counted as two valves. The

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species relative abundance was expressed in number of valves per 2 gr of dry residue.

3.2 Benthic foraminifers

Benthic foraminifers were randomly picked from the fraction greater than 0.125 mm.

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In most cases, more than 200 specimens were picked per sample. The specimens were

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identified according to Cimerman and Langer (1991) and Milker and Schmiedl (2012).

3.3 Invertebrates (mollusks) The bulk samples were gently wet-sieved through standard screens to enable the picking of fossil invertebrates. This technique yielded a number of relatively well-preserved skeletal parts ranging in size from several centimeters down to the microfossil grade. When the carbonate matrix was not broken down satisfactorily by natural weathering, the skeletal material was mechanically extracted and included in the study. Specimens from all fractions above 0.25 mm grain size were picked, whereas the fractions below 0.25 mm were not investigated further. The abundance of skeletal material was specified semi-quantitatively,

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ACCEPTED MANUSCRIPT considering shell orientation, disarticulation, fragmentation, and encrustations to distinguish whether or not the shells had been transported before their final deposition.

3.4 Calcareous nannofossils Smear slides for calcareous nannofossil biostratigraphic analysis were prepared following the standard preparation technique of Perch-Nielsen (1985) and Bown and Young (1998). To obtain accurate biostratigraphic estimations, up to 100 fields of view were

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investigated per slide. All samples were routinely examined at 1,250× using a LEICA DMLSP light microscope (LM). The total assemblage composition was evaluated in a count,

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where all specimens (coccoliths/nannoliths) were counted until a statistically significant total,

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of at least 500 nannofossils, was obtained. The semi-quantitative abundances of the taxa encountered were recorded as follows: abundant (more than one specimen in every field of

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view), common (one specimen in 10 fields), rare (one specimen in 50 fields), present (one specimen in more than 100 fields), and reworked specimens. We followed the biozonation of Martini (1971) and the biochronological framework of Lourens et al. (2004), Raffi et al.

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(2006), and Backman et al. (2012); all these schemes provided biohorizon ages for the eastern

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Mediterranean.

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4. Results

4.1. Akropotamos sections

The Akropotamos Neogene deposits are severely disturbed by normal faulting;

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therefore, a long continuous section is unavailable. Four sections (A, B, C, and D) are studied, which are located in three abandoned gypsum quarries aligned the first three in a SW–NE

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direction, whereas section D in a WNW–ESE direction (Fig. 5). Despite the major problems imposed by the Neogene sequence fragmentation, stratigraphic correlations are made using two indexes: 1) the transition between the uppermost part of the gypsum beds and the overlying laminites with terrestrial plant fragments, and 2) a massive carbonate bed in the upper part of the sections (indicated by the light blue shade in Fig. 5).

4.1.1. Section A It is located in the escarpment of the western gypsum quarry (Figs. 5 and 6 A). The lower 6.5 meters of the sequence consist of gypsarenites and interbedded massive microcrystalline gypsum lenses topped by one meter calcareous gypsarenites. The upper 2.5 8

ACCEPTED MANUSCRIPT meters are composed of alternating decimeter-thick interbeds of silty limestones and laminites.

4.1.2 Section B Section B is located between the western and the median quarry, about 30 meters east of Section A (Figs. 5 and 6 B), from which it is separated by a normal fault. It is the only section in the study area where the uppermost part of the pre-evaporitic sequence can be

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observed. It is exposed below the evaporitic unit, after a 5-meter observation gap, for about 4.5 meters. The pre-evaporitic sequence consists of clastic deposits marked by marine marl

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intercalations with bivalves fragments (pectinids, oysters), passing to an open marine facies at

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the top. Fossils are either absent or rare to very rare; the most common are plant leaves (molds), fish remains (mostly bone fragments and scales), and ostracods. They are sometimes

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accompanied by benthic foraminifers, serpulid worms, bivalves, gastropods, echinoids (spines only), bryozoans, and siliceous sponge spicules.

The evaporitic sequence begins with 1.80 m argillaceous and carbonate laminites

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(Figs. 5 and 6 B), followed by a 2.3-meter-thick succession of interbedded massive microcrystalline gypsum (0.70 m), sandy gypsarenite (0.70 m), and interbedded massive

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microcrystalline gypsum (0.90 m). The following 3.5-meter-thick succession comprises: 1.2 m argillaceous/carbonate laminites; 0.40 m green marls with bivalves, serpulids (Fig. 6 D),

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and coal; 0.60 m limestones with rare bivalves; approximately 0.10 m marls; 0.50 m

4.1.3 Section C

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limestones with plants; and 1.20 m dark marls.

This section is located in the escarpment of the median quarry, about 600 m northeast

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of Sections A and B (Figs. 5 and 6 C). The lower 6 m consist of interbedded massive microcrystalline gypsum (2.5 m), followed by 2 m alternating argillaceous and carbonate laminites, and topped by 1.5 m interbedded massive microcrystalline gypsum (Fig. 6 C). The following 13-meter-thick upper part of the section begins with 3 m alternating laminites and laminated limestones, where fossil leaves are observed. The remaining 10 m of the sequence mainly consist of laminites, with fish remains in the middle part of the succession, intercalated by rare thin silty limestone beds. The top of the laminites is intercalated by 0.80 m dark clays followed by 0.70 m massive limestones.

4.1.4 Section D 9

ACCEPTED MANUSCRIPT This section is located in the escarpment of the eastern quarry, about 600–650 m northeast of the median quarry’s Section C (Figs. 5, 6, and 7). The lower 7 m begin with 1 m interbedded massive microcrystalline gypsum; followed by 1 m gypsum limestone with fenestrae, in which a microbialite layer of 2–3 mm thickness is observed (sample 2b; Fig. 6 G), and clayey carbonates with serpulids; 3 m alternation of metric to decametric gypsrudite to gypsarenite, calcareous gypsum (Fig. 6 H), limestones with interbedded massive microcrystalline gypsum nodules, and laminated limestones with resedimented gypsum beds.

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The unit ends with 2 m laminites and sandy limestones with serpulids and bivalves, as well as rare gypsum lenses. The uppermost part of the gypsum unit is capped by an angular

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unconformity, since the strata of the overlying sequence onlap the upper surface of the

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gypsum unit (yellow line; Fig. 7). The overlying part of the section (post-evaporitic sequence) is accessible for another 13 meters. It consists of a succession beginning with 1 m laminated

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limestones with convolute structures and argillaceous laminites, followed by 9 m laminites. The laminites display some silty clay intervals with ripple marks and some silt to limestone interbeds with hummocky cross-stratification. In their lower part, the laminites also yield

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leaves, some fish fragments, and bryozoans. They are followed by some 0.60-m-thick dark clays, then by 1.5 m porous fine-grained limestone, and 2 m laminated marls with 0.40 m of

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ferruginous limestone at their base. In the uppermost part of Section D, limestone thin sections display high porosity, relictual fenestral fabric, pisoids, brecciation, and diffuse

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peloidal micrite invaded by iron hydroxides. Such features indicate lacustrine deposits and possible emergence events. The unit is capped by a 3-m travertine layer (Fig. 7). This carbonate layer displays millimeter-sized plant fragments embedded in a cloudy micritic

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matrix with locally cauliflower stromatolitic microstructures. It is noteworthy that the travertine constitutes a marker horizon observed in many outcrops correlated across the entire

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Strymon Basin. This observation, first made by Snel et al. (2006b), is crucial for the interpretation of the environmental conditions during travertine deposition in the entire Strymon Basin.

The contact between the highest travertine bed and the overlying clastic marine deposits of the Strymon Basin is not exposed in the studied sections (Fig. 7). Nevertheless, the wider area consists of predominantly clastic (conglomerate, sandstone, clay) deposits, which can reach a maximum thickness of 1500 m across the Strymon Basin. The open marine clastics and marl alternations at the base of this sequence were previously dated in the Zanclean (Steffens et al., 1979; Karistineos and Georgiades-Dikeoulia, 1986; Snel et al., 2006b; Suc et al., 2015). 10

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4.1.5 Gypsum facies Gypsum facies correspond mainly to massive, microcrystalline gypsum (fine-grained, snow-white homogeneous gypsum, known also as alabaster or saccharoidal gypsum; Figs. 6 B and E), and secondary to gypsarenite/gypsrudite intercalations (Fig. 6 F). Microcrystalline gypsum is created, where anhydrite pervasively rewaters the gypsum in the zone of active phreatic flow. This can be caused when evaporite beds are exhumed and uplifted and thus

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entering the telogenetic realm (Warren et al., 1990). Microcrystalline gypsum and gypsarenites have undergone a cycle of dehydration–hydration, whereas the different textures

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displayed by these facies can be regarded as different stages of progressive anhydrite

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hydration (Testa and Lugli, 2000). Αnhydrite represents the precursor of microcrystalline gypsum, and could be interpreted as a diagenetic product, derived from dehydration of a

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primary gypsum deposit (Murray, 1964; Holliday, 1970). Gypsrudite/gypsarenite layers of Akropotamos sections, which occasionaly include some larger gypsum clasts in the form of lenses, could originate from the reworking of in situ cumulate gypsum or derived from the

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erosion and transport of the broader Strymon area gypsum deposits. Microbialite deposits are considered as a product of the interactions between activity

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of benthic microorganisms and physiochemical properties of environment. The millimeterthick microbialite layer intercalated in the gypsum limestone of the Akropotamos section D

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(sample 2b; Figs. 5 and 6 G) probably correspond to continuously rising water (Bąbel, 2005). Activity of currents is one of the most important physical processes influencing the accretion of microbialite domes. The development of microbial mats, found directly below

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microcrystalline gypsum, probably represent a very shallow ephemeral pan deposited under permanent brine. This event requires periodic fluctuations in salinity at the beginning of the

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gypsum saturation stage. Salinity falls favored the development of specific mat-forming benthic microbial communities (Bąbel, 1999, 2004, 2005; Bąbel et al., 2011, 2015).

4.2 Paleontological findings 4.2.1 Ostracods, benthic foraminifers, and invertebrates The Akropotamos sections bear an ostracod fauna of 32 species referable to 23 genera. Some characteristic species are illustrated in Fig. 8. The qualitative and semi-quantitative ostracod analysis indicates differences in the composition of the ostracod assemblages along the studied sections (Fig. 9).

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ACCEPTED MANUSCRIPT The preservation of benthic foraminifers is generally moderate to poor in most samples. Reworked specimens with filled tests, abraded surface, and a red patina of hydrous iron oxide are present in many samples, particularly in those with a higher percentage of sand. The most abundant species are Valvulineria complanata, Rosalina globularis, Cibicides lobatulus, and Ammonia beccarii. Their relative abundances vary throughout the studied sections and, overall, they suggest reworking. Porosononion granosum occurs in relatively high abundance only in sample CK12A. The ‘high-productivity/low-oxygen’ species group is

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dominated by Bolivina spp., Brizalina spp., and Uvigerina peregrina, and it is only found in sample BK’18.

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The macrosamples contain a relatively diverse mollusk fauna, composed of bivalves

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and a few gastropods, accompanied by annelids and echinoids. Several bivalves and gastropods with decalcified shells have been collected; many bivalves show articulated

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valves. In some cases, only proximally thickened shell parts or fragments of thin shells are available. The gastropods in the limestones are found as molds, but, in laminites, they are well

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preserved.

4.2.2 Ostracod assemblages and biostratigraphic implications

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Ostracod assemblages at levels AK11, BK14, BK12, BK'1, BK'4, BK'7, and CK12A are oligotypic and marked by the significant presence of Cyprideis agrigentina and

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Loxoconcha muelleri (Fig. 9). C. agrigentina is a Mediterranean euryhaline species found in the Lago Mare deposits all over the Paleo-Mediterranean (Ligios and Gliozzi, 2012; Grossi et al., 2015). L. muelleri is a Paratethyan immigrant, also distributed in the Mediterranean

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during the post-evaporitic Messinian (Faranda et al., 2007). Ostracod assemblages of very low diversity, dominated by C. agrigentina and L. muelleri, are common during the lower

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Lago Mare interval (Gliozzi et al. 2007; Trenkwalder et al., 2008; Faranda et al., 2013), forming the L. muelleri zone (5.59–5.40 Ma; Grossi et al., 2011). Therefore, the horizons corresponding to the above samples are placed within this zone. These two species are also dominant in AK11, BK12, and CK12A, where they are accompanied by few individuals of Heliocythere? anura (AK11) or Dorukella bireticulata (CK12A). However, their state of preservation in these samples indicates transportation. In samples BK14, BK'1, BK'4, and BK'7, they are the most abundant species and accompanied by several species, among them Amnicythere cf. A. propinqua. Amnicythere is a Paratethyan leptocytherid, which entered the Mediterranean during the Lago Mare event, and it is present with several species in the post-evaporitic Messinian ostracod assemblages (e.g., Gliozzi et 12

ACCEPTED MANUSCRIPT al., 2005). A. propinqua has been found in several Messinian Lago Mare deposits (Italy, Gliozzi et al., 2005; Crete, Cosentino et al., 2007; Spain, Guerra-Merchán et al., 2010). Dorukella bireticulata (sample BK14) has been described from the Tortonian (Darbaş et al., 2008) and the Messinian (Doruk, 1974) sediments of Turkey, as well as the late Miocene of Skyros Island (Guernet, 2005). Accordingly, Semicytherura sanmarinensis (sample BK’4) is known from the early Tortonian of Crete (Faranda et al., 2008) and the early Messinian of Italy (Ruggieri, 1967), at the onset of the MSC (Italy, Gennari et al., 2013).

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Both species are present with very few individuals.

Concerning samples BK'5 and BK'8, the ostracod assemblages are composed of

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several species similar to those found in BK14, BK'1, BK'4, and BK'7, including L. muelleri,

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but in low abundances. The same applies for sample BK'21, where L. muelleri is not present but Loxocorniculina sp. is, a Paratethyan immigrant that was spread into the Mediterranean

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during the Lago Mare event (Krstic', 1972; Faranda et al., 2007). Significant faunal alternations take place from sample CK11 downward. Sample CK11 includes the most diversified and abundant ostracod fauna, composed mainly of

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Loxocorniculina sp., Palmoconcha agilis, Euxinocythere (Maeotocythere) cf. E. (M.) praebaquana, C. excanaliculata, Xestoleberis species, A. convexa accompanied by L. ovulata,

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Callistocythere producta, C. agrigentina, C. acuminata, Cytherois sp., D. bireticulata, Paracytheridea sp., and Aurila spp. Besides Loxocorniculina sp., the presence of the genus

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Euxinocythere in CK11 indicates Paratethyan influence, as it is a Paratethyan species which migrated into the Mediterranean during the final stage of the MSC (Gliozzi, 1999; Gliozzi et al., 2002). Consequently, sample CK11 could be attributed to the post-evaporitic Messinian.

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C. excanaliculata was also recorded in the early Messinian of Skyros Island, Greece (Bonaduce and Russo, 1985; Bonaduce et al., 1992; Babinot and Boukli-Hacene, 1998).

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However, in Akropotamos sections, it is present in several samples with low frequencies (BK14, BK11, BK’4, BK’21; poorly preserved indicating transportation), except for sample CK11, where it is present in great numbers. Thus, its stratigraphic range could be expanded. Likewise, C. producta known from the early Messinian (Italy, Aruta, 1982; Algeria; Babinot and Boukli-Hacene, 1998) is also present in this diversified assemblage. Samples CK10 and CK12B contain an oligotypic assemblage, where Heliocythere? anura is the dominant species. Heliocythere? anura has been described from the late Messinian deposits of Tunisia (Bonaduce et al., 1992), and it is an enigmatic species in the Akropotamos ostracod assemblages. It is present in sample AK11, but with evidence of transportation, and it forms nearly monospecific assemblages in CK10 and CK12B, which is 13

ACCEPTED MANUSCRIPT above CK12A and the C. agrigentina-L.muelleri assemblage. Likewise, Limnocythere sp. is dominant in the assemblages of samples CK5 and CK4. Finally, the character of the ostracod assemblage changes once again in CK2B. Loxocauda aff. L. decipiens is the dominant species, present in great numbers and accompanied by Xestoleberis species, Paradoxostoma sp., P. turbida, C. excanaliculata, Cytherois sp., D. bireticulata, Loxoconcha sp.3, and Semicytherura sp. L. decipiens and P. turbida have a wide stratigraphic range, therefore the presence of D. bireticulata and C.

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excanaliculata points to a Messinian age.

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4.2.3 Calcareous nannoplankton

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Only Sections C and D yield significant calcareous nannofossil assemblages. In Section C, the studied samples are barren of calcareous nannofossils, except for samples

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BK’7 and BK’21. Sample BK’4 contains aragonite spicules. Sample BK’7 reveals the poorest assemblage with very rare and extremely small coccoliths of Syracosphaera spp. and very rare specimens of Lithostromation perdurum. Sample BK’21 consists of a very rich

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assemblage of Helicosphaera spp. (abundant H. carteri, common–rare H. sellii) and Syracosphaera spp. Mostly S. pulchra, but also Coronosphaera spp. are recorded; specimens

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of L. perdurum are abundant.

In Section D, the studied samples are barren of calcareous nannofossils, except for

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CK11, which contains abundant Syracosphaera spp. and H. carteri, as well as common–rare H. sellii and L. perdurum. In addition, Discoaster spp. are common, including Discoaster bergrenii. Rare specimens of Amaurolithus primus are identified; Nicklithus amplificus is

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absent. The contemporaneous presence, in samples CK 11 (Section D) and BK’21 (Section C), of Discoaster bergrenii and Amaurolithus primus supports the biostratigraphic assignment

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of the studied samples to biozone NN11, in particular NN11b (7.424–5.54 Ma; Raffi et al., 2003, 2006; Krijgsman et al., 2004; Backman et al., 2012; Fig. 5). Notably, Discoaster bergrenii, Amaurolithus primus, and Helicosphaera sellii, which are found here in the upper part of the evaporitic unit, were also recorded at the same stratigraphic position in other localities of the Strymon Basin (Snel et al., 2006b).

4.3 Akropotamos paleoecologic implications The paleoenvironmental evolution in the Akropotamos area is reconstructed based on the analysis of the ostracod, benthic foraminifer, and invertebrate assemblages recovered from the clastic deposits intercalated in the gypsum unit and from the overlying Lago Mare 14

ACCEPTED MANUSCRIPT deposits. Most samples point to a coastal sheltered lagoonal environment with salinity fluctuations due to the communication with the sea and/or freshwater input. The identified macrofauna, microfauna, and nannoflora indicate that, in the lower part of the Akropotamos sections, the gypsum limestones and laminated marls were deposited in waters with fluctuating salinities, ranging from shallow-marine to oligohaline. Their sedimentary structures with microbial mats, ripple marks, and dominant parallel-bedding lamination indicate a low-energy depositional environment. The Akropotamos sequence overlying the

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gypsum unit corresponds to a lagoon episodically linked to the open sea. The uppermost parts of the sections, with limestones, reflect lacustrine depositional environments.

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In particular, we distinguish eight stages in the paleoenvironmental evolution of the

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Akropotamos area at 5.9–5.4 Ma: four stages between 5.9 and 5.6 Ma (MES), in the interval corresponding to the gypsum unit; and four stages between 5.6 and 5.4 Ma (L. muelleri zone

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above MES), in the lower part of the Lago Mare unit. The paleoecological results are summarized in Figure 10.

Stage 1. The ostracod assemblages in the basal part of section D (samples CK2A and

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CK2B) are composed mainly of Loxocauda aff. L. decipiens, which is a shallow marine taxon (Tsourou, 2012), and it is accompanied mainly by Xestoleberis spp., Paradoxostoma sp., P.

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turbida, Callistocythere excanaliculata, Cytherois sp., and Semicytherura sp. This assemblage is indicative of a shallow marine infralittoral environment (Clave et al., 2001; Bracone et al.,

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2012) with subaquatic vegetation, as Xestoleberis spp. and Paradoxostoma sp. are highly associated with algae (Athersuch, 1976; Cronin et al., 2001; Triantaphyllou et al., 2005). Additionally, some Xestoleberis species tolerate significant salinity fluctuations (Mazzini et

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al., 1999; Cronin et al., 2001; Viehberg et al., 2008). The most abundant benthic foraminifer species are Valvulineria complanata, Rosalina globularis, Cibicides lobatulus, and Ammonia

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beccarii, and they are, though badly preserved, indicative of an inner shelf environment (0– 100 m water depth). Likewise, Serpula spp. reefs, small ostreids, and rare Chlamys multistriata indicate shallow marine high-energy conditions. Stage 2. The dominance of the ostracod taxa Limnocythere sp. (samples CK4 and CK5) indicates a freshwater to oligohaline environment. In sample CK4, fragments of Hydroides (serpulids) and ostreids, as well as the presence of many Ditrupa fragments suggests marine influence. Stage 3. This stage is characterized by an oligotypic ostracod assemblage (sample CK10) dominated by Heliocythere? anura, which is considered a marine species (Seko et al.,

15

ACCEPTED MANUSCRIPT 2015), and it is accompanied by the euryhaline Xestoleberis spp. Consequently, this assemblage reflects a polyhaline to euhaline shallow marine environment. Stage 4. The assemblages in this stage (samples BK′21 and CK11) correspond to a similar paleoenvironment that is a sheltered coastal polyhaline lagoon. In particular, sample BK′21 bears a mixed ostracod assemblage with oligo-mesohaline taxa such as Loxocorniculina (Faranda et al., 2007), Limnocythere, and Cyprideis, which are accompanied by shallow infralittoral species (Xestoleberis spp., A. convexa, and L. ovulata), pointing to a

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mesohaline to polyhaline waterbody. The benthic foraminiferal assemblage is nearly monospecific with Valvulineria complanata, indicating a rather sheltered environment with

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salinity variations. The predominance of Valvulineria complanata further supports the great

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availability of organic matter, mainly brought by river runoff and leading to a poorly oxygenated sea floor (van der Zwaan and Jorissen, 1991). The mollusk fauna is characterized

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by Apomatus sp. (forming rather loose aggregates floating in the sediment), pectinid fragments, Anomia ephippium, Anodontia fragilis, Tellina sp., and Plagiocardium, which point to a shallow marine environment. However, the fauna also includes Mactra sp.

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(brackish) and Mytilopsis cf. frici (freshwater). The unattached free ends of Apomatus sp. indicate a low-energy environment such as a lagoon. The mixing of the fauna points probably

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to a coastal lake, which was periodically invaded by seawater. Likewise, a mixed ostracod assemblage occurs in sample CK11, with brackish Paratethyan ostracod taxa, such as

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Loxocorniculina and E. (M.) praebaquana (low mesohaline and shallow waterbodies; Grossi et al., 2015); the euryhaline C. agrigentina; and several shallow marine infralittoral species with abundant P. agilis (shallow coastal marine species; Boomer et al., 2010), C.

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excanaliculata, Xestoleberis spp., A. convexa, L. ovulata, C. acuminata (epineritic species; Seko et al., 2012), and C. producta. The foraminiferal fauna is characterized by the presence

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of C. lobatulus, C. refulgens, A. beccarii, Rosalina carnivora, Bulimina aculeata, Gutulina communis, Pyrgo spp., Quinqueloculina spp., Elphidium complanatum, B. gibba, and Discorbis mira. The co-occurrence of foraminifers living in different environmental settings seems to indicate that these specimens are transported from nearby environments. The freshwater hydrobiids Pseudamnicola, which are grazers and deposit feeders (Gofas, 2010), occur in slowly running spring-fed freshwater bodies and channels (Djamali et al., 2006), but also in slightly brackish estuaries, and even in saline spring streams (Moreno et al., 2010) along with Mytilopsis sp. (freshwater mollusk), small ostreids, Anomia ephippium, echinoids spines, Anodontia fragilis, Tellina sp, Plagiocardium, Mactra sp., and fragmented Serpula spp., probably indicating a coastal lake invaded by seawater. 16

ACCEPTED MANUSCRIPT Stage 5. The ostracod fauna in BK11, composed of Xestoleberis spp., A. convexa, Loxoconcha ovulata, Paradoxostoma sp., and Callistocythere spp., indicates a shallow marine infralittoral environment (Bracone et al., 2012; Clave et al., 2001). Xestoleberis spp. and Paradoxostoma sp. as well as L. ovulata (Zangger and Malz, 1989; Lachenal, 1989) suggest a shallow marine environment rich in algae. The benthic foraminiferal fauna indicates increased turbidity, consistent with enhanced rainfall and fluvial runoff. In particular, in sample BK11, the predominance of Valvulineria complanata indicates a rather sheltered environment with

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salinity variations and a poorly oxygenated sea floor. The mollusk fauna in BK11 indicates a high-energy marine infralittoral environment with fragmented serpulid buildups of Hydroides,

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thin shelled pectinids, and many specimens of Anomia ephippium.

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Stage 6. The monospecific ostracod assemblage of sample BK12, with C. agrigentina but in low numbers, is attributed to a high mesohaline shallow-water (10–15 m) environment

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(Grossi et al., 2008). The mollusks in BK12 form a mixed fauna of Gastrochaenolithes, Scala (Striatiscala?) sp., Ostrea sp., Corbula sp., Tellina sp., Abra sp., Venus sp., Mactra cf. faugeresi (Paratethyan species), Pirenella sp., Sphaeronassa? sp., Polinices sp., and

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Dentalium sp., characteristic of shallow marine to lagoon environments. Pirenella is an euryhaline species that inhabits coastal and inland lagoons on fine, sandy, or muddy substrates and also occurs in saline and hypohaline lakes in deltas, where it can tolerate

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extreme temperatures (5–45°C) and salinities (15–90 ‰), although it cannot survive in

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habitats exposed to waves (Taraschewski and Paperna, 1981). Therefore, its abundance in sample BK12 points to an open lagoon environment. Stage 7. The fossil findings in this stage (samples AK11, BK14, BK′7–BK′1, and

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CK12A) point to a low mesohaline lagoonal environment with seawater influence, no more than 10–15m deep, with short-term dysoxic conditions. The ostracod assemblages are

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oligotypic and they are composed mainly of abundant C. agrigentina and L. muelleri, which indicate a low mesohaline environment, no more than 10–15-m deep (Grossi et al., 2008). In samples BK14 and BK′7–BK′1, these species are accompanied by the freshwater taxon Limnocythere, the oligo-mesohaline Amnicythere cf. A. propinqua (Guerra-Merchán et al., 2010) and Xestoleberis spp. There are also marine specimens present in BK14 as A. convexa, Callistocythere excanaliculata, and D. bireticulata but they may be reworked. Benthic foraminiferal fauna is present in sample BK14, but it is reworked and characterized by a mixture of epifaunal and infaunal taxa of the infralittoral realm. In particular, the benthic foraminiferal fauna is characterized by Cibicides lobatulus, Cibicidoides italicus, Bigenerina nodosaria (few), Hanazawaia boueana, Elphidium fichetellianum, E. macellum, Bulimina 17

ACCEPTED MANUSCRIPT aculeata, B. subulata, Cassidulina laevigata, Bolivina spathulata-dilatata, B. pseudoplicata, and Uvigerina spp., all indicating a coastal marine environment. In samples BK′7–BK′1, benthic foraminifers are badly preserved suggesting reworking. On the other hand, the benthic foraminiferal fauna of CK12A is characterized by the presence of Porosononion granosum, Ammonia beccarii, and Bolivina seminuda, which form an euryhaline assemblage, mostly typical of lagoonal-paralic environments without vegetation, at mid- to low-tidal elevations. The mollusk fauna of BK14, BK’4, and CK12A is characterized by juvenile thin-shelled

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sessile bivalves. Rare echinoid spines are also present. Within a lagoon depositional setting, small bivalves live as endo-byssate non-siphonate suspension feeders. These bivalves are

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interpreted as juveniles colonizing firm, dysoxic carbonate substrates. Short-term dysoxic

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phases interpreted to reflect seasonal decreases in water energy resulted in the mass mortality of juvenile bivalves (Bassi et al., 2015).

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Stage 8. Finally, the uppermost sample with micropaleontological content is CK12B (Section D). It comprises H. anura, C. agrigentina, and Leptocythere sp., probably indicating a mesohaline to polyhaline waterbody. Benthic foraminifers are characterized by the presence

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of Bolivina seminuda, Rosalina, and Elphidium, indicating an inner shelf environment (0–100 m water depth). The high frequency of Bolivina seminuda suggests high productivity/low

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oxygen conditions.

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5. Discussion

5.1 Stratigraphy and sedimentology

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5.1.1 Onshore facies integression during the MSC stages In order to establish the age of the onshore Akropotamos evaporites and their

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correlation with the offshore Prinos-Nestos evaporitic unit, it is very important first to locate the Messinian erosional surface (MES) in both onshore and offshore sequences. Indications of the MES in the Strymon Basin are reported by Suc et al. (2015) that attributed the unconformity east of Serres City (Karistineos and Georgiades-Dikeoulia, 1986) to an erosional surface between the evaporitic beds and the overlying clastic deposits, interpreting the latter as foreset beds of Gilbert-type fan delta deposits. In general, in the peripheral Mediterranean basins, the MES has been placed at 5.60 Ma (Roveri et al., 2014a). In the Ionian Basin (Zakynthos Island), it has also been assigned the same age (Karakitsios et al., 2017). In the present study of the Akropotamos sections, the ostracod associations at the base of the deposits overlying the evaporitic unit indicate an age 18

ACCEPTED MANUSCRIPT interval 5.59–5.42 Ma for this stratigraphic level, whereas the nannofossil association observed in the upper part of the evaporites indicate an upper limit 5.54 Ma (zone NN11b; 7.424–5.54 Ma). Based on these ages, there are two possible positions of the MES: a) in the upper part of the evaporitic unit, and b) at the base of the evaporitic unit. In the first case, the angular unconformity observed on the top of the evaporitic unit (yellow lines; Fig. 7) can be correlated with the MES marking the passage to the post-evaporitic unit (CIESM, 2008; Roveri et al., 2014a). The non-observation of the erosional unconformity in the other sections

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(A, B, C) is due to the fact that the quarries exposing these sections are aligned in a SW–NE direction, which is parallel to the direction of the strata, and this constitutes the only case

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where it is impossible to observe an existing unconformity. Thus, in this case, the MES

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merges with the surface at the base of the Lago Mare deposits. This scenario is also in agreement with the dominance of microcrystalline gypsum facies in the gypsum unit, since

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this facies is considered a diagenetic product derived from the dehydration of a primary gypsum deposit (Murray, 1964; Holliday, 1970). Besides, the intercalated microbialite layer in the Akropotamos section D (Figs. 5 and 6 G), directly below the microcrystalline gypsum,

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probably represents a very shallow ephemeral pan deposited under permanent brine, requiring periodic fluctuations in salinity at the beginning of the gypsum saturation stage that favors the

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development of benthic microbial mat (Bąbel, 1999, 2004, 2005; Bąbel et al., 2011). The second case, according which the MES is located as a correlative conformity at the base of the

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evaporitic unit, and the evaporitic unit corresponding to RLG deposits, is excluded by the observations that were reported in the first case. Besides, the attribution of the gypsum unit to the RLG deposits is also contrary to the fact that RLG deposits are formed in depths greater

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than 300 m (from the slope area to the depocenter of the evaporitic basin; Lugli et al., 2015). Consequently, we conclude that, in Akropotamos, the MES is located between the base of the

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Lago Mare and the top of the gypsum unit, which could be attributed to the MSC stage 1 (PLG; 5.97–5.6 Ma; Roveri et al., 2014a; Fig. 5). The entire shale and limestone sequence overlying the gypsum unit, represents brackish environments with variable salinity similar to the Paratethyan depositional environments (as indicated by the ostracod, mollusk, and nannofloral assemblages).The age interval 5.59–5.42 Ma of its base could attribute the entire sequence to the Lago Mare unit, deposited during MSC stage 3 (5.5–5.33 Ma; Riding et al., 1998; Krijgsman et al., 1999; Roveri et al., 2014a; Lugli et al., 2015; Fig. 5). The travertine bed above this sequence indicates a freshwater depositional environment. Based on its stratigraphic position, between the shallow marine gypsum unit and the more open marine Pliocene clastic deposits, the 19

ACCEPTED MANUSCRIPT travertine probably represents the local continental expression of the Lago Mare biofacies. In the younger marine clastic deposits overlying the travertine beds, close to the Akropotamos village, calcareous nannofossils belonging to Zone NN12, and indicating an early Pliocene age were reported by Steffens et al. (1979).

5.1.2 Offshore facies integression during the MSC stages and onshore-offshore correlations Offshore, in the Nestos slope area of the Prinos-Nestos Basin (Fig. 4), the benthic

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foraminifer Uvigerina rutila is continuously present 60 m above the evaporitic unit of the Nestos-2 well (Sidiropoulos, 1980). This species is a widely used biostratigraphic marker of

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the lower part of the Mediterranean Pliocene and supports a Zanclean age for the base of the

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post-evaporitic unit (Iaccarino, 1985; Suc et al., 2015). In addition, the presence of the nannofossil Discoaster quinqueramus in the uppermost part of the evaporitic unit (40 m

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below its top) in Nestos-2 well (Sidiropoulos, 1980) gives an upper age limit of 5.54 Ma (Raffi et al., 2006) for the corresponding stratigraphic level. This suggests that the entire evaporitic unit could be older than 5.6 Ma. Thus, considering that the base of the post-

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evaporitic sequence corresponds to 5.33 Ma, which is the lower limit of the Zanclean, the 60 m inbetween could correspond to the Lago Mare. Thus, the MES should be an unconformity

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positioned above the evaporitic unit (anhydrite), which then should be assigned to the PLG (5.6–5.97 Ma).

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In an attempt to find the prolongation of the MES in the Prinos deep area, we examine the published SW–NE seismic profile across the Prinos field (Proedrou and Sidiropoulos, 1992; Proedrou and Papakonstantinou, 2004; Fig. 11). In this profile, we observe that, in the

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left half of the profile, the reflection terminations of the post-evaporitic sequence are subparallel to those of the evaporitic unit (halite), as well as to the boundary between them,

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whereas, in the right half of the profile, they onlap the previous strata and boundary, showing evidence of erosion, as is deduced by the truncation of underlying salt unit reflections. Thus, the left half of the profile could be related to the Top Surface (TS) marker, whereas the right half to the Top Erosional Surface (TES) marker. TS and TES correspond to markers determined by offshore seismic analysis (Lofi et al., 2011a; both TS and TES were previously referred by Ryan (1973) as Younger M surface). In the offshore places of the Mediterranean, TS represents the basinward conformable surface, whereas when erosion indications are present the TES represents the slopeward erosional surface (Lofi et al., 2011b). From this point of view, the entire salt unit could represent the offshore Mobile Unit (MU) composed by halite and clastics (MU, Lofi et al., 2011a, b), which is considered to be deposited during the 20

ACCEPTED MANUSCRIPT 5.97–5.33 Ma interval (Roveri et al., 2014a). Alternatively, the salt unit and the anhydrite clastics observed at the base of the Prinos salt unit (Fig. 4), which are more abundant toward the Nestos area (Pollak, 1979), combined with the truncation of the upper halite intercalation toward the Nestos slope area, suggest that the lower part of the salt unit (bottom clastic products included) could be deposited during the acme of the MSC that was reached during the second stage (RLG stage 2; 5.6–5.55 Ma; Roveri et al., 2014a), whereas the upper part of the salt unit could represent the equivalent of the Upper gypsum (UG, 5.55–5.33; Roveri et

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al., 2014a). Τhis interpretation appears to be better supported by the existing data. Concerning the two salt intercalations of the Prinos well, predating the MSC (Fig. 4),

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they could be attributed to the early Messinian preparatory stage of the MSC (7.251–5.5.97

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Ma; Roveri et al., 2014a). This stage corresponds to the gradual restriction of the marine connections to the Atlantic, and it is commonly related to tectonic uplift processes in the

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gateway area (Duggen et al., 2003; Garcia-Castellanos and Villaseñor, 2011), whereas the first evidence of significant restriction in Mediterranean-Atlantic exchange was recorded by reduction of deep-water ventilation all over the Mediterranean at 7.15 Ma immediately after

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the Tortonian-Messinian boundary (Kouwenhoven et al., 1999, 2003; Seidenkrantz et al.,

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2000).

5.1.3 The halite-shale couplets depositional environment

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So far, the data from both the western and eastern deep Mediterranean basins only refer to the uppermost evaporite units. Indeed, according to Lugli et al. (2015), the Messinian giant evaporite deposits of the Mediterranean were sampled only for several meters by a few

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cores drilled in the 70s and 90s (decision taken to avoid a possible hydrocarbon blowout in the Mediterranean). These units mainly consist of clastic (gypsrudite, gypsarenite, and

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gypsiltite) and fully subaqueous deposits (laminar gypsum, selenite, and cumulate halite) that are partially affected by burial anhydritization and tectonically induced recrystallization (Hardie and Lowenstein, 2004; Lugli et al., 2015). Ηence, Lugli et al. (2015) concluded that the above sedimentological characters do not constitute unequivocal evidence of shallow water or even supratidal (sabkha) deposition. It was suggested that, at the very last phase of the MSC, the Mediterranean Sea did not experience desiccation, but deposition took place under permanent subaqueous conditions (Roveri et al., 2014b; Lugli et al., 2015; Carnevale et al., 2017). In the Prinos deepest area, the comformable strata between the post-evaporitic and the evaporitic sequence (deduced by the reflection terminations), and the marine shales’ presence 21

ACCEPTED MANUSCRIPT immediately above the evaporitic unit (halite-shale couplets) as well as within the shale intercalations of the evaporitic unit show that, despite the sea drawdown, halite accumulation took place under permanent marine conditions during the MSC (Figs. 4, 11 and 12). The rare case of the offshore Prinos-Nestos Basin, where the entire evaporitic sequence has been totally penetrated by boreholes in the depocenter and the slopes, shows that the deep-water deep-basin model (Schmalz, 1969, 1991; De Benedetti, 1976, 1982; Dietz and Woodhouse, 1988; Roveri et al., 2014b) could provide an explanation for the deep salt deposition. The

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cyclical alternance of clastic sediments supply and halite is considered to be, as in the case of gypsum-shale couplets, directly induced by astronomically driven climatic changes with a

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precession periodicity (Krijgsman et al., 1999; Krijgsman and Meijers, 2008; Lugli et al.,

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2010; Roveri et al., 2014a).

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5.2 The Mediterranean-Paratethys connection issue

Snel et al. (2006a) indicated several marine ingressions in the Dacic Basin during the late Miocene and the early Pliocene, based on marine nannofossil assemblages.The Paratethys

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became connected to the marine realm at 6.12 Ma (Krijgsman et al., 2010; Stoica et al., 2013; Chang et al., 2014; van Baak et al., 2016). A connection to the Mediterranean has also been

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indicated throughout the MSC stage 1, based on hydrogen isotope analyses performed in the Black Sea (Vasiliev et al., 2013; 2015), bringing Paratethyan water to the Mediterranean and

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possibly contributing to the PLG formation (Natalicchio et al., 2014). Furthermore, Suc et al. (2015) hypothesized that, during the MSC peak, a major drainage system was developed along the Struma-Strymon corridor, bringing significant clastic material into the northern

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Aegean; this hypothesis supports the model that freshwater input contributed to evaporite deposition during the MSC (Natalicchio et al., 2014). A connection was present during the

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final MSC stage, leading to the deposition of the Lago Mare facies, which comprises typical Paratethyan fauna (Gliozzi, 1999; Stoica et al., 2016). There are several possibilities for the location of the Mediterranean-Paratethys gateway(s): A.

North Aegean–Black Sea (Dardanelles/Sea of Marmara)

A gateway was indeed established in this region during the Pontian (Steiniger & Rogl, 1984; Popov et al., 2006; Krijgsman et al., 2010). However, van Baak et al. (2016) supported that fully marine conditions were not established in the Sea of Marmara between the Serravallian and the late Pliocene, citing the paleogeographic study of Cagatay et al. (2006), which however, does not exclude periodic connections in this area. 22

ACCEPTED MANUSCRIPT B.

North Aegean–Dacian Basin (Strymon Basin)

A gateway was proposed along the Strymon-Struma Basin by Snel et al. (2006b) and Suc et al. (2015). However, van Baak et al. (2016) do not support this option, basing their argument on the study of Spassov et al. (2006), which however refers to some localities along the Struma River, but not to the entire river area. It is therefore, entirely possible that these continental environments correspond to the banks of the gateway. C.

Eastern Mediterranean–Black Sea (through SE Turkey)

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Van Baak et al. (2016) indicated that several mostly freshwater subbasins existed throughout SE Turkey during the late Miocene containing faunas of Paratethyan affinity (e.g., Denizli

Eastern Mediterranean–Caspian Sea (through Zagros)

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D.

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basin; Wesselingh et al., 2008). However, their connectivity still requires investigation.

Van Baak et al. (2016) also postulated the existence of small subbasins connecting the eastern

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Mediterranean to the Caspian Sea. Indeed, a significant sea-level drop that would preclude a connection between the Capsian Sea and the Black Sea, also dismissing the unification of the Paratethys basins, is not considered possible based on recent hydrologic models (de la Vara et

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al., 2016). However, evidence has only been presented for the Tabriz Basin in northern Iran (Reichenbacher et al., 2011).

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The Paratethys may have been connected to the Indian Ocean at 6.12 Ma, a possibility that would explain the presence of radiolarians and foraminifera, including Streptochilus that

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only inhabited the western Indian Ocean, in the eastern Paratethys during this flooding event (Krijgsman et al., 2010; Stoica et al., 2013; van Baak et al., 2016). However, at least a oneway gateway to the Mediterranean would be necessary to allow the Paratethyan fauna to enter

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the Mediterranean Basin during the Lago Mare phase. This Paratethyan water influx to the Mediterranean would be observable during the Lago Mare phase, if there was Atlantic water

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influx to the Mediterranean, even not continuously, as has been previously suggested (Grunert et al., 2016; Cornee et al., 2016), which would raise the sea level enough for the Paratethyan fauna to migrate to the Mediterranean. Such an assumption is supported by and compatible with the presence of marine fish within the Lago Mare facies (Carnevale et al., 2006; 2008; 2017), and it is in agreement with previous observations in the northeastern Aegean (Sakinc & Yaltirak, 2005) as well as the recent alkenone data from Sicily and the Black Sea (Vasiliev et al., 2017), which in fact indicate a Mediterranean-Black Sea connection during this time as well. In fact, both Sakinc & Yaltirak (2005) and Cagatay et al. (2006) report alternations of Mactra-bearing and Ostrea-bearing limestones in the late Messinian of the Bosporus/Sea of Marmara area/Gulf of Saros, which correspond to freshwater and marine conditions, 23

ACCEPTED MANUSCRIPT respectively. It is possible that the freshwater deposits were deposited during Mediterranean Sea lowstands, when Paratethyan water outflow was greater toward the Aegean Sea, through the Sea of Marmara. On the other hand, during Mediterranean Sea highstands, the Aegean Sea was reconnected with the Mediterranean, allowing for the establishment of marine conditions in the study area. The Mediterranean-Atlantic connection is the key to the expression of this phenomenon, because this would be necessary for the re-introduction of the marine fish in the western Mediterranean. The temporary nature of these Mediterranean-Atlantic connections

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during the final MSC stage is also justified by the absence of other marine organisms beside fish, such as foraminifera and marine mollusks, which would require a longer time interval

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and more pervasive marine conditions in order to re-inhabit the Mediterranean Basin.

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The results presented here indicate that the study area was a coastal sheltered lagoonal environment with salinity fluctuations due to the communication with the sea and freshwater

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input during the MSC stage 1, that corresponds to the PLG deposition. These observations support that the northern Aegean Sea was in communication with the rest of the Mediterranean Sea during this stage, but also that it received considerable freshwater and

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faunal input from the Paratethys, as indicated by the mixed marine-brackish fauna, which intensified stratification contributing to the evaporite deposition, as suggested by Natalicchio

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et al. (2014). Above the proposed MES, the Lago Mare sequence was deposited under both marine and freshwater influence. Therefore, we conclude that the study area remained

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connected to the Mediterranean and the Paratethys throughout the MSC, even if not continuously. The uppermost part of the sequence in this area is characterized by the travertine deposition, which indicates that, at the end of the Lago Mare deposition, just before

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the Pliocene reflooding, the study area in the northern Aegean, including Akropotamos and

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probably the upper part of Nestos-Strymon slope, was a freshwater lake.

5.3 Depositional model and paleogeographic interpretation The onshore biostratigraphic data from the Strymon Basin show that it constituted a peripheral basin where the PLG unit was deposited. An erosional phase is observed by the MES on top of this unit. Above the MES, there are brackish-water Lago Mare deposits, which represent the final stage of the MSC. The Paratethyan fauna in these deposits has fueled a long-lasting controversy over the connectivity between the Mediterranean and the Paratethys (e.g., Psilovikos and Syrides, 1983; Snel et al., 2006b; Suc et al., 2015), and the contemporaneous sea-level drop in both basins has been attributed to the isolation of the Mediterranean from the Atlantic Ocean (Clauzon et al., 2005; Popov et al., 2006; Popescu et 24

ACCEPTED MANUSCRIPT al., 2009; Suc at al., 2011; Bache et al., 2012; Krstić et al., 2012; Bakrač et al., 2012; Do Couto et al., 2014; Suc et al., 2015). In the Akropotamos area, during the Lago Mare deposition, the infilling of the basin could have led to the formation of coastal lagoons with fluctuating salinities progressively changing upward into lakes. This could have been achieved through the seaward progradation of the Lago Mare during a still-stand or a limited sea-level drop. Thus, during the Lago Mare deposition, the sea drawdown, in combination with the subsidence of the area, was insufficient to expose the area to erosion, but a balance

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between Paratethyan and Mediterranean waters was installed, as it is recorded in the alternation between freshwater and marine conditions, respectively. Toward the end of the

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Lago Mare deposition, when the sea-drawdown reached its maximum, the sea level was found

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below the level of the Paratethyan waters causing the input of freshwater over the area, creating lakes where travertines were deposited on top of the Lago Mare facies. Consequently,

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the travertine bed in the Strymon Basin refers to the upper part of the Lago Mare unit (Fig. 13). The wide lateral continuation of this marker bed throughout the Strymon Basin (Snel et al., 2006b) suggests that the corresponding lacustrine environment was extensive over the

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entire basin. The above considerations are in accordance with the results of Vasiliev et al. (2017), according to which the similarity between δDalkenones recorded during the MSC stage 3

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from Mediterranean sites and the Paratethys shows that surface water during Upper Gypsum deposition must be derived from Paratethyan water inflow into the Mediterranean. Thus, the

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Prinos-Nestos Basin Lago Mare deposits corresponded first to lagoons with changing salinities and then to freshwater environments. However, evidence for a prolonged phase of erosion persisting until the early Pliocene

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in the marginal basins of the Mediterranean Sea (Clauzon et al., 1996) is not found here. Instead, predating the open marine Pliocene deposits, in the Akropotamos sections, we

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observe the sequence: evaporite–brackish water deposits–travertine beds. This sequence exhibits a wide lateral continuation throughout the Strymon Basin. In addition, marine nannofossils were previously recorded in the Pliocene marls overlying the travertine beds, in other sections of the Strymon Basin (Steffens et al., 1979; Snel et al., 2006b). These data suggest that marine normal conditions were restored after an erosional phase, which took place during the sea drawdown before the deposition of Lago Mare. In previous studies, the marine ingressions inferred in the deep eastern Mediterranean basins (in the marginal basins of the North Aegean) and in the Dacic Basin, before the beginning of the Pliocene, were used to question the Mediterranean deep basin-shallow water scenario for the MSC (Castradori, 1998; Mǎrunţeanu and Papaianopol, 1998; Snel et al., 2006a). The presence of brackish-water 25

ACCEPTED MANUSCRIPT faunas and nannofossil assemblages in the highest part of the Messinian (see subsection 4.2.3 and Fig. 10) indicates that, before the early Pliocene, the basin was not completely desiccated. In this part of the Northern Aegean, the Miocene–Pliocene transition was mainly marked by a salinity change, caused by the increasing interaction between Atlantic-Mediterranean and Paratethyan waters, predating the marine reflooding attested by the following Pliocene marine deposits. According to Cornée et al. (2016) and to the model of Marzocchi et al. (2016), the end of the MSC was not caused by a catastrophic flooding at the Miocene–Pliocene boundary,

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but by the reorganization of the circulation patterns and the establishment of MediterraneanAtlantic water exchange similar to today.

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Offshore, in the adjacent Prinos-Nestos Basin, the evaporite facies and thickness

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distribution indicate that, during the upper Messinian sea drawdown, PLG deposits accumulated in the slope areas. During the maximum sea drawdown, the erosional phase of

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the slope PLG fed the deeper part of the basin with gypsum clastics, whereas the halite saturation in the restricted deep area during the peak of the MSC caused also its accumulation, while the PLG deposits were being eroded in the marginal/peripheral basin (slope area).

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Because of the very expressed cyclicity of the seven halite-shale couplets, it is difficult to accept that heavy brines and erosional products from the slope evaporitic unit were reached seven times by downslope flows of hypersaline, dense waters, in a process similar to present

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day “dense shelf water cascading” (Roveri et al., 2014b; Lugli et al., 2015). This case could

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only be discussed for the lowest and thickest halite layer. In this case, the downslope flows of the hypersaline dense waters, facilitated by the the marginal basin’s faults and the slope’s sliding surfaces, could accumulate the heavy brines (salt) in the deeper part of the basin over

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resedimented gypsum and clastics. However, we consider as more probable that the total cyclical alternance of halite and shale is directly induced, as in the case of gypsum-shale

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couplets, by astronomically driven climatic changes with the precession periodicity (Krijgsman et al., 1999; Krijgsman and Mejiers, 2008; Lugli et al., 2010; Roveri et al., 2014a), and the halite was always deposited in the deep part of the basin, despite the sea drawdown, under relatively deep seawater, as the intercalated and overlying marine shales suggest. In an attempt to classify the studied onshore-offshore basins of the area, we consider that the Strymon (Akropotamos) Basin could be ascribed to the shallow marginal basins, the Nestos Basin to the intermediate marginal basins, and the Prinos Basin to the intermediate to deep basins (Roveri et al., 2014a). Thus, in this relatively small area of study, the main

26

ACCEPTED MANUSCRIPT evaporitic depositional facies of the Mediterranean are represented, as well as their relationship with the Paratethys.

6. Conclusions Our results on the late Neogene onshore Strymon and its correlation with the offshore Prinos-Nestos deposits lead to the following conclusions (Fig. 13). In the Akropotamos area, the MES is located on top of the evaporitic unit, which has

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been deposited during the MSC stage 1 (5.97–5.6 Ma). Over the Messinan erosional surface was deposited the Lago Mare unit during the MSC stage 3 (5.55–5.33 Ma). This final stage of

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the MSC in Akropotamos area was characterized by brackish-water deposits that end by a

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travertine bar, which the extensive presence in the entire onshore Strymon Basin, indicates that small lagoons and lakes were probably established along the Mediterranean-Paratethyan

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corridor predating the high sea-level phases. The Miocene–Pliocene transition in the Akropotamos area was marked by a salinity change caused by the increasing influence of Paratethyan over Atlantic-Mediterranean waters, predating the marine reflooding at the end of

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the MSC, which is attested by the open marine Pliocene Gilbert-type delta deposits overlying the evaporites.

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The evaporitic unit of the offshore Prinos-Nestos sequence was deposited during the upper Messinian Sea drawdown phase in a marine environment. In the slope areas of the

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basin, the evaporitic unit is composed of anhydrite-clastic/shale couplets, passing to a thick sequence of halite-shale couplets in the deeper part of the basin. The first sediments (lower Pliocene) of the post-evaporitic sequence were deposited in a marine environment, which

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became progressively more open marine. The evaporites (anhydrites) of the Nestos slope area were deposited in the MSC stage 1 (PLG 5.97–5.6 Ma), whereas the evaporites (salt) in the

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Prinos deepest area were deposited under permanent marine conditions, probably during the MSC stage 2 and 3 (5.6–5.33 Ma).

Acknowledgments This research has been co-financed by the European Union (ESF) and Greek national funds through the Operational Program "Education and Lifelong Learning" of the National Strategic Reference Framework (NSRF) - Research Funding Program: THALIS –UOA-“Messinian Salinity Crisis: the greatest Mediterranean environmental perturbation and its repercussions to the biota” MIS: 375405. We are grateful to the two anonymous reviewers, whose constructive comments greatly improved the manuscript. 27

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Planet. Sc. Lett. 362, 272–282. http://dx.doi.org/10.1016/j.epsl.2012.11.038.nzi. Vasiliev, I., Reichart, G.-J., Grothe, A., Sinninghe Damsté, J.S., Krijgsman, W., Sangiorgi, F., Weijers, J.W.H., van Roij, L., 2015. Recurrent phases of drought in the upper Miocene of the Black Sea region. Palaeogeogr. Palaeoclimatol. Palaeoecol. 423, 18–31. http://dx.doi.org/10.1016/j.palaeo.2015.01.020 Vasiliev, I., Mezger, E.M., Lugli, S., Reichart, G-J., Manzi, V., Roveri, M., 2017. How dry was the Mediterranean during the Messinian Salinity Crisis? Palaeogeography, Palaeoclimatology, Palaeoecology, doi: 10.1016/j.palaeo.2017.01.032. Viehberg, F.A., Frenzel, P., Hoffmann, G., 2008. Succession of late Pleistocene and Holocene ostracode assemblages in a transgressive environment: A study at a coastal locality of 41

ACCEPTED MANUSCRIPT the southern Baltic Sea (Germany). Palaeogeography, Palaeoclimatology, Palaeoecology 264(3–4), 318–329. Warren, J. K., Havholm, K. G., Rosen, M. R., Parsley, M. J., 1990. Evolution of gypsum karst in the Kirschberg Evaporite Member near Fredericksburg, Texa. Journal of Sedimentary Petrology, 60, 721–734. Wesselingh, F.P., Alcicek, H., Magyar, I., 2008. A Late Miocene Paratethyan mollusc fauna

palaeobiogeographic implications. Geobios 41, 861–879.

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from the Denizli Basin (southwestern Anatolia, Turkey) and its regional

Wüthrich, E.D., 2009. Low Temperature Thermochronology of the Northern Aegean

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Rhodope Massif. PhD thesis (ETH No. 18673), Swiss Federal Institute of Technology

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Zurich, 210 pp.

Zangger, E., Malz, H., 1989. Late Pleistocene, Holocene and Recent Ostracods from the Gulf

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of Argos, Greece. Courier Forschungsinstitut Senckenberg 113, 159–175.

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ACCEPTED MANUSCRIPT Figure captions

Fig. 1. Simplified paleogeographic map showing the depositional facies distribution in the Mediterranean-Paratethys region during the late Messinian (modified from Rouchy and Caruso, 2006).

Fig. 2. Simplified geological map of the greater area containing the Akropotamos onshore and

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the Prinos-Nestos offshore basins (modified after Kydonakis et al., 2015). Core complexes cropping out in the area: Northern Rhodope Domain (NRD), Southern Rhodope Core

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Complex (SRCC), Chalkidiki block, Vardar s.l. units, and Pelagonia. The term Northern

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Rhodope Core Complex (NRCC) is used collectively for the exhumed gneiss domes (Chepinska, Arda, Kardamos-Kesebir, and Kechros-Biela-Reka domes) exposed to the

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northeast within the NRD, whereas the term Southern Rhodope Core Complex (SRCC) refers to the triangle-shaped gneiss dome exposed at the central part of the map. The rectangle frame

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at the center of the map corresponds to the study area (Fig. 3A).

Fig. 3. A) Simplified geological map of the Akropotamos–Prinos-Nestos area. Gray circles

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synthetic cross section of Fig. 12. Green markers correspond to boreholes.

Fig. 4. Representative columns of the offshore Prinos and Nestos areas (based on Lalechos

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and Savoyat, 1979; Proedrou, 1979; Proedrou and Sidiropoulos, 1992). In the Nestos column are positioned the characteristic fossils determining the ages of the top of the evaporitic unit,

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and the base of the post-evaporitic unit. In the Prinos column the marine shales (with typical marine fossil association) frame the Messinian salt unit and are also present in the marl intercalations of this unit. In the same column note the anhydrite clastics at the base of the salt unit.

Fig. 5. Akropotamos sections lithostratigraphic correlation and sampling. MES line indicates the possible position of the Messinian erosional surface.

Fig. 6. Sedimentary facies of the Akropotamos sections: A: interbedded massive microcrystalline gypsum and gypsarenite sequence followed by alternating decimeter-thick 43

ACCEPTED MANUSCRIPT interbeds of silty limestones and laminites (Section A), B: argillaceous and carbonate laminites (0.80 m), followed by a 2.3 meter-thick succession of interbedded massive microcrystalline gypsum (0.70 m), sandy gypsarenite (0.70 m) and stratified interbedded massive microcrystalline gypsum (0.90 m) (Section B), C: stratified interbedded massive microcrystalline gypsum alternations (red arrows) into argillaceous and carbonate laminites (section C), D: marls with serpulids (Section B), E: interbedded massive microcrystalline gypsum, F: resedimented gypsum elements included into marls, G: microbialite layer of 2–3

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gypsum.

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Fig. 7. I: Panorama of Section D. The red rectangle frame corresponds to the detail shown in J; the red line on top of the exposed deposits marks the boundary of the travertine layer with

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the underlying sequence. J: The red line shows the erosional unconformity, above which overlap the strata of the Lago Mare unit (yellow lines).

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Fig. 8. Scanning electron microscope pictures of selected ostracod species from the Akropotamos sections. Lateral external views (RV: right valve, LV: left valve) of some of the

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most abundant ostracod species in the studied samples: 1-3. Cyprideis agrigentina Decima: 1. Female, LV, sample AK 11; 2. Female, RV, sample AK 11; 3. Male, LV, sample CK 4. 4, 5.

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Loxoconcha muelleri (Mehés), sample AK 11: 4. Female, LV; 5. Female, RV. 6. Callistocythere excanaliculata Bonaduce and Russo, LV, sample CK 2B. 7. Callistocythere producta Aruta, RV, sample CK 11. 8. Cytheridea acuminata (Bosquet), LV, sample CK 4. 9.

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Aurila convexa (Baird), LV, sample BK’4. 10. Heliocythere anura Bonaduce et al., LV, sample CK 10. 11. Dorukella bireticulata (Doruk), RV, sample CK 11. 12. Pontocythere

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turbida (Müller), RV, sample CK 2B.

Fig. 9. Distribution and frequency of the identified ostracod taxa in the studied samples and the determined age along the sections.

Fig. 10. A concise and comparative presentation of the paleoecological results.

Fig. 11. Northeast-southwest interpreted seismic section across the Prinos field (for location see Fig. 3B where the wells referred in the seismic section are positioned) (modified after Proedrou and Sidiropoulos, 1992 and Proedrou and Papaconstantinou, 2004). A SW-dipping 44

ACCEPTED MANUSCRIPT fault that formed the Prinos field roll-over anticline and separates it from the North Prinos structure merges with a sole fault that underlies the evaporitic section in the North Prinos area. The graben between Prinos and North Prinos is the result of a NE-dipping antithetic fault. Pre-Neogene Alpine basement dips to the SW. Boreholes Prinos-2, Prinos-4, and Prinos-6 appear in Fig. 3 as P-2, P-4, and P-6, respectively. “Top of pay” refers to the top of the productive reservoir, which, in this case, is immediately below the base of the evaporitic

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unit. The numbers on the right vertical axis correspond to sec.

Fig. 12. Synthetic cross section along the long axis of the Prinos-Nestos Basin (based on

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borehole and seismic data from Lalechos and Savoyat, 1977; Pollak, 1979; Proedrou, 1979,

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1988; Sidiropoulos, 1980; Proedrou and Sidiropoulos, 1992; Proedrou and Papaconstantinou, 2004; Harker, 2008). The Prinos field reservoirs are located immediately below the main

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evaporitic unit, whereas in the South Kavala field the reservoir is within the evaporitic unit. We consider that the MES unconformities (raffled line) at the basin’s margins (Nestos) pass to a correlative conformity (continuous line) at the base of the evaporitic unit in the deeper

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part of the basin (Prinos). Boreholes: South Kavala (SK4 and SK2), Prinos (P1), Ammodhis (A1), Nestos (N2 and N1); s.l.: sea level. The position of the section is shown in the Fig. 3

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Fig. 13 Simplified paleogeographic evolution of the study areas. A. MSC stage 1. PLG deposition takes place in the Akropotamos shallow-marine basin and in the Nestos intermediate basin. Deposition in the deep Prinos Basin includes marine shales and

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carbonates. The entire area is connected to the Mediterranean and receives freshwater input from the Paratethys. The Paratethyan faunal content in Akropotamos increases upward.

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B. MSC acme. Aerial erosion of the PLG deposits takes place in the Akropotamos area, while subaerial to subaqueous erosion affects the Nestos Basin. The Prinos basin receives anhydrite clastics originating from the adjacent Nestos and maybe even Akropotamos areas. C. MSC stage 3. Lago Mare is deposited in the Akropotamos and Nestos basins, while the halite-shale couplets are formed in the Prinos Basin. At the end of this stage, the Akropotamos area is completely isolated from the marine environment, is dominated by freshwater input from the Paratethys, and sees the deposition of the travertine bed. light blue: freshwater, dark blue: seawater. The undulating part and the continuous part of the red line correspond to the unconformity and the correlative conformity of the MES, respectively.

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low mesohaline environment, no more than 10–15m deep, a lagoon with sea water influence and short-term dysoxic facies

Biostratigraphy

Loxoconcha muelleri zone Lower part of Lago Mare event (Grossi et al., 2011)

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CK12B Ostracods: H. anura, C. agrigentina, Leptocythere sp. Foraminifera: Bolivina seminuda, Rosalina, Elphidium CK12A Ostracods: C. agrigentina , L. muelleri Foraminifera:Porosononion granosum, Ammonia beccarii, and Bolivina seminuda Mollusks: juvenile thinshelled sessile bivalves

Environmental interpretation mesohaline to polyhaline lagoon, high productivity/low oxygen conditions

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BK14 Ostracods: C. agrigentina, L. muelleri, Limnocythere sp. Amnicythere, Xestoleberis spp. Mollusks: juvenile thin-shelled sessile bivalves BK12 Ostracods: C. agrigentina Mollusks: Gastrochaenolithes, Scala (Striatiscala?) sp. Ostrea sp., Corbula sp., Tellina sp., Abra sp., Venus sp., Mactra cf. faugeresi, Pirenella sp., Sphaeronassa? sp., Polinices sp., Dentalium sp. BK11 Ostracods: Xestoleberis spp., A. convexa, Loxoconcha ovulata, Paradoxostoma sp., Callistocythere spp Foraminifera: predominance of Valvulineria complanata. Mollusks: fragmented serpulid buildups of Hydroides, thin shelled pectinids, Anomia ephippium

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AK11 Ostracods: C. agrigentina, L. muelleri

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a shallow marine infralittoral environment with increased turbidity, consistent with enhanced rainfall and fluvial runoff

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Paleoenv. stages 8

BK′21 mixed assemblages Ostracods: Loxocorniculina, Limnocythere, Cyprideis, Xestoleberis spp., A. convexa, and L. ovulata) Foraminifera: predominance of Valvulineria complanata. Mollusks: Apomatus sp., Anomia ephippium, Anodontia fragilis, Tellina sp., Plagiocardium, Mactra sp. Mytilopsis cf. frici.

MES CK11 mixed assemblages Ostracods: Loxocorniculina, E. (M.) praebaquana, C. agrigentina; P. agilis, C. excanaliculata, Xestoleberis spp., A. convexa, L. ovulata, C. acuminata, C. producta. Foraminifera: C. lobatulus, C. refulgens, A. beccarii, Rosalina carnivora, Bulimina aculeata, Gutulina communis, Pyrgo spp., Quinqueloculina spp., Elphidium complanatum, B. gibba, and Discorbis mira. Mollusks: Pseudamnicola, Mytilopsis sp., small ostreids, Anomia ephippium, echinoids spines, Anodontia fragilis, Tellina sp, Plagiocardium, Mactra sp., fragmented Serpula spp. CK10 Ostracods: Heliocythere? anura, Xestoleberis spp. CK4, CK5 Ostracods: Limnocythere sp. Mollusks in CK4: fragments of Hydroides,

A sheltered coastal polyhaline lagoon

Discoaster bergrenii and Amaurolithus primus biozone NN11b (Raffi et al., 2006; Backman et al., 2012)

polyhaline to euhaline shallow marine environment freshwater to oligohaline environment marine influence in

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CK4 shallow marine infralittoral environment with subaquatic vegetation. Highenergy conditions

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