ORIGINAL PAPER

Checklist of benthic foraminifera (class Foraminifera: d’Orbigny1826; phylum Granuloreticulosa) from Saros Bay, northernAegean Sea: a biodiversity hotspot

Fabrizio Frontalini & Michael A. Kaminski &Ioulietta Mikellidou & Eric Armynot du Châtelet

Received: 9 December 2013 /Revised: 2 May 2014 /Accepted: 2 May 2014# Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2014

Abstract Calcareous and agglutinated benthic foraminiferaldiversity was investigated in the central Northern Aegean Sea.A total of 18 samples were collected along a neritic to mid-bathyal transect in Saros Bay at water depths ranging from 15to 500 m. This study reveals a very highly diversified benthicforaminiferal fauna composed of 267 species belonging to 115genera. Benthic foraminiferal taxa varied along the depthtransect, and it is possible to recognise benthic foraminiferalassemblages typical for different bathymetric ranges. An in-ner–middle neritic assemblages (ca. <100 m depth), outerneritic assemblages (ca. 100–200 m), and upper bathyal

assemblages (200–500 m) are defined. On the basis of theavailable foraminiferal literature on the Aegean Sea coupledwith our data from Saros Bay, a very high number of forami-niferal species is estimated to occur in this basin. A total of799 species belonging to 276 genera, 93 families and 43superfamilies have been recorded in the Aegean Sea. Thesevalues are remarkably higher than those documented in pre-vious foraminiferal studies carried out in different parts of theMediterranean Sea. Although the determination of the totalforaminiferal diversity in the Mediterranean Sea remains adifficult task, we speculate that this number may be in excessof 1,000.

Keywords Benthic foraminifera . Checklist . Saros Bay .

Aegean Sea

Introduction

The Rhizaria represents one of the six supergroups of eukary-otes based on molecular data and phylogenetic analyses (Cav-alier-Smith 2002; Nikolaev et al. 2004; Sierra et al. 2013) andincludes various flagellate and amoeboid protists such as theforaminifera. Historically, the study of foraminifera has beenthe domain of palaeontologists, and specifically applied topetroleum exploration and to palaeoclimatological andpalaeoecological reconstructions. More recently, the study oftheir recent living counterparts has kindled the interests offoraminiferologists, particularly for biodiversity, biological,ecological and biomonitoring studies (Murray 2006). Forami-nifera represent one the most abundant and diversified shelledtaxa in our oceans (Sen Gupta 1999). More than 2,000 recentliving benthic foraminiferal species have been reported, andthe overall worldwide diversity is estimated to be around3,000–4,000 (Murray 2007). Numerous studies have beenpublished on benthic foraminifera from different marine

Electronic supplementary material The online version of this article(doi:10.1007/s12526-014-0238-z) contains supplementary material,which is available to authorized users.

F. Frontalini (*)Dipartimento di Scienze della Terra, della Vita e dell’Ambiente(DiSTeVA), Università degli Studi “Carlo Bo”, Campus Scientifico,Località Crocicchia, 61029 Urbino, Italye-mail: fabrizio.frontalini@uniurb.it

M. A. KaminskiDepartment of Earth Sciences, King Fahd University of Petroleumand Minerals (KFUPM), Dhahran, Saudi Arabiae-mail: kaminski@kfupm.edu.sa

M. A. Kaminski : I. MikellidouDepartment of Earth Sciences, UCL, Gower Street, London WC1E6BT, UK

I. MikellidouRPS Energy, Guildford, UKe-mail: mikellidoui@rpsgroup.com

E. Armynot du ChâteletUniversité Lille 1, UMR 8217 CNRS Géosystèmes,UFR Sciences de la Terre, Bât SN5, Avenue Paul Langevin,59655 Villeneuve d’Ascq, Francee-mail: eric.armynot@univ-lille1.fr

Mar BiodivDOI 10.1007/s12526-014-0238-z

basins such as the Mediterranean Sea, but most of them dealwith areas of limited geographical extent, and mainly the near-shore environments. The Mediterranean Sea represents a clas-sic locality to investigate foraminiferal diversity (i.e., Parkerand Jones 1860; Sidebottom 1904, 1905, 1906, 1907, 1908,1909; De Rijk et al. 1999, 2000; Milker and Schmiedl 2012),and detailed taxonomical studies have been carried out indifferent parts of it. In particular, important contributions havebeen provided by Lacroix (1928, 1929, 1930, 1932a, b, 1933,1935), Todd (1958), Bizon and Bizon (1984), Schmiedl et al.(2000), Fontanier et al. (2008), Mojtahid et al. (2009),Goineau et al. (2011), Contreras-Rosales et al. (2012) andMilker and Schmiedl (2012) for the western Mediterranean,Parker (1958) and Cimerman and Langer (1991) for theeastern part, Jorissen (1988), De Stigter et al. (1998), andDonnici and Serandrei Barbero (2002) for the Adriatic Sea,Alavi (1988), Kaminski et al. (2002), Phipps et al. (2010) andFrontalini et al. (2011) for the Marmara Sea; Buchner (1940),Hofker (1960) and Sgarrella and Moncharmont Zei (1993) forthe Gulf of Naples (western Italy), Basso and Spezzaferri(2000) for Iskenderun bay (Turkey), and Meriç et al. (2008a)for the Antalya coast, among others.

Although several investigations have been carried out inthe Aegean Sea area, to our knowledge no comprehensivestudy has documented the benthic foraminiferal diversity anddistribution along a bathymetric gradient from the shelf todeep basin. Following this, the aim of this paper is to docu-ment the modern benthic foraminiferal fauna and their bathy-metric distributions along an inner neritic to mid-bathyaltransect from 15 to 500 m water depth in Saros Bay and, bycompiling data from the previous investigations, to provide anestimate of the benthic foraminiferal diversity in the AegeanSea. It must be underlined that the Aegean Sea represents akey area in term of hydrological mechanisms being the placewhere Eastern Mediterranean Deep Water (EMDW) isformed. Moreover, alien and invasive species related to globalwarming might strongly impact the biodiversity balance inthis area in the next years/decades along with substantialconsequences for its marine ecosystems.

Study area, geography and hydrology

The Aegean Sea, an elongated embayment occupying one ofthe northern extremities of the Eastern Mediterranean Sea, isplaced between the southern Balkan and Anatolian peninsulasand, by the island of Crete in the south and Rhodes, Karpathosand Kasos to the southeast (Fig. 1). The central and north-western part of the Aegean area is mainly formed by thenumerous islands and archipelagos that include Cyclades,Ikaria, Samos, Dodecanese, and Sporades. The general circu-lation within the Aegean Sea is cyclonic with high salinemarine water inflowing from the Levantine basin further south

and travelling northwards along the western coast of Turkey(Theocharis et al. 1999). The Aegean Sea is also stronglycharacterised by freshwater discharges of small rivers andseasonal inflows of modified Black Sea water through theDardanelles straits. This inflow influences the uppermost(down to 40 m) layer of the North Aegean Sea (Theocharisand Georgopoulos 1993; Pazi 2008), with its strongest influ-ence in Saros Bay. This water mass is then modified movingwestward and southward by mixing with the Levantine Inter-mediate Water (Stergiou et al. 1997). Although the AegeanSea has been considered as an oligotrophic environment,significant differences exist between the relatively nutrientenriched north–northwest and the less nutrient enrichedsouth–southeast parts of the Aegean Sea (Stergiou et al.1997). Accordingly, the outflow of the Black Sea into SarosBay leads to an enrichment of dissolved organic carbon anddissolved organic nitrogen (Polat and Tuğrul 1995, 1996),whereas the southern part of the Aegean Sea has been consid-ered as one of the most oligotrophic areas of the Mediterra-nean Sea (i.e., Pazi 2008 and references therein). Furthermore,it must be stressed that the Aegean Sea is a strategic area interm of hydrological mechanisms representing one of theplace where EMDW is formed. The formation of EMDWcan be regarded as an important component of the Mediterra-nean thermohaline circulation (Rubino and Hainbucher 2007).The same authors documented a major transformation in thethermohaline cell of the eastern Mediterranean Sea where theAdriatic Sea has returned to represent a major source ofEMDW after two decades. They pointed out, however, thatthe characteristics of such water profoundly differ from thosepreviously observed before and during the Eastern Mediterra-nean Transient (EMT) being saltier, warmer, and denser.

Saros Bay, also known as the Gulf of Saros, is an inlet inthe northeastern part of the Aegean, north of the GallipoliPeninsula of northwestern Turkey (Figs. 1, 2). The waterdepth reaches ca. 700 m, and continues towards the WSW tojoin the 1,000- to 1,500-m-deep North Anatolian Trough.Meriç River in the northwest and Kavak Creek in the eastare the main sources of fresh water and sediments for SarosBay (Sarı and Çağatay 2001).

Previous benthic foraminiferal studies in the Aegean Sea

Although several papers have documented the benthic fora-miniferal distribution in selected parts of the Aegean Sea,most studies have only focused on selected areas of this basin,and several of them deal with fossil foraminiferal assemblages(i.e., Rasmussen 2005; Abu-Zied et al. 2008; Drinia et al.2004; Drinia and Anastasakis 2012). To our knowledge, thefirst benthic foraminiferal investigations in the Aegean Seawere carried out on Delos island (1; numbers in parenthesesrefer to Fig. 1) by Sidebottom (1904, 1905, 1906, 1907, 1908,

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Fig. 1 The Aegean Sea showing the investigated areas. 1 Delos island:Sidebottom (1904, 1905, 1906, 1907, 1908, 1909); 2 Çeşme-Ilica Bay:Avşar and Meriç (2001), Bergin et al. (2006), Meriç et al. (2012a) andBarut et al. (2013); 3 Gokceada Island: Avşar and Ergin (2001), Avşar(2002), Meriç et al. (2001a, 2004b), Yalçın et al. (2008) and Bassler-Veitet al. (2013); 4 Bozcaada Island: Avşar and Ergin (2001), Meriç et al.(2001a, 2004b), Avşar (2002) and Yalçın et al. (2008); 5 Canakkale:Avşar and Ergin (2001), Avşar (2002) and Meriç et al. (2004b); 6 Gulfof Saros: Meriç et al. (2002b, 2004a, b); 7 the Gulf of Edremit (Meriçet al. 2003, 2004b) and Bassler-Veit et al. (2013); 8 central Crete: Driniaet al. (2004); 9 Kallithea Bay: Rasmussen (2005); 10 Saros Gulf: Meriç(1996), Meriç et al. (2001b, 2004b, 2005, 2008b), Bassler-Veit et al.(2013); 11 near the island of Bozcaada: Meriç (1996), Meriç et al.(2001b, 2004b, 2005, 2008b) and Yalçın et al. (2008); 12 the EdremitGulf: Meriç (1996) and Meriç et al. (2001b, 2004b, 2005, 2008b); 13Gökova Gulf: Meriç (1996) and Meriç et al. (2001b, 2004b, 2005,2008b); 14 Korthi and Kastro gulfs (Andros Island) Triantaphyllouet al. (2005, 2009, 2012) and Koukousioura et al. (2011); 15 Gulf ofKalloni (Lesvos Island, Greece): Debenay et al. (2005); 16 easternMitilini: by Yalçın et al. (2008); 17 Pamucak Cove: (Meriç et al. 2008c,

2009a, 2010a); 18 Büyük Menderes River delta: Avşar et al. (2009); 19Dardanelles: Meriç et al. (2009b); 20 Falasarna: Triantaphyllou et al.(2009, 2012); 21 Porto Heli: Triantaphyllou et al. (2009, 2012); 22MavroLithari: Triantaphyllou et al. (2009, 2012); 23 Vravron: Triantaphyllouet al. (2009, 2012) and Koukousioura et al. (2012); 24 Kalamos:Triantaphyllou et al. (2009, 2012); 25 Katerini: Triantaphyllou et al.(2009, 2012); 26 N. Kallikratia: Triantaphyllou et al. (2009, 2012); 27Avdira: Triantaphyllou et al. (2009, 2012); 28 Alykes Kitros:Koukousioura et al. (2012); 29 Lafrouda Thrace: Meriç et al. (2004b),Koukousioura et al. (2012), Bassler-Veit et al. (2013) and Frontalini et al.(2014); 30, 31 Gulf of Saros: Meriç et al. (2004b), Bassler-Veit et al.(2013); 32 Biga Peninsula: Bassler-Veit et al. (2013); 33 Canakkale:Meriç et al. (2004b); 34 Gulf of Edremit: Meriç et al. (2004b); 35 DikiliBay: Meriç et al. (2004b); 36 Candarli Bay: Meriç et al. (2004b); 37Karaburun Peninsula: Meriç et al. (2004b, 2012c, d); 38 Gulf ofKusadasi: Meriç et al. (2004b); 39 Gulluk Bay: Meriç et al. (2004b); 40Gulf of Datca: Meriç et al. (2004b); 41 Marmaris Bay: Meriç et al.(2004b); 42 Illica Bay: Meriç et al. (2012b, c, d); 43 Camalti saltpan:Meriç et al. (2010b)

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1909) (Fig. 1). The distribution of recent benthic foraminiferafrom a thermal region in Çeşme-Ilica Bay (Izmir) was reportedby Avşar and Meriç (2001) (2) who listed a total of 40calcareous foraminiferal species including Coscinospirahemprichii, Laevipeneroplis karreri, Peneroplis pertusus,Peneroplis planatus and Sorites orbiculus (Fig. 1). The distri-bution of Holocene benthic foraminifera in the Gokçeada-Bozçaada-Canakkale (3–5) was investigated by Avşar andErgin (2001) and Avşar (2002) who documented the presenceof 160 benthic foraminiferal species belonging to 84 genera(Fig. 1). Later, Meriç et al. (2001a) recognised 104 speciesbelonging to 58 genera near Gokçeada Island (3), a slightlylower diversity was documented in Bozcaada Island (4) where58 species belonging to 26 genera were identified (Meriç et al.2002a). In Saros Bay (6), a quite well-diversified benthicforaminiferal assemblage including, in total, 163 species be-longing to 95 genera were recognised (Meriç et al. 2002b,2004a). In the Gulf of Edremit (7), 101 benthic foraminiferalspecies were listed by Meriç et al. (2003). Diversity andabundance trends of late Pliocene benthic foraminifera fromthe southern part of central Crete (8) were investigated byDrinia et al. (2004). In the Plio-Pleistocene Kallithea Bay(Rhodes, Greece southern–eastern Aegean Sea), a total of249 taxa of benthic foraminifera were recognised by Rasmus-sen (2005) (9). The presence of some abnormal nodosariidsand other benthic foraminiferal species were documented inSaros Bay (10), near the island of Bozcaada (11), in theEdremit Gulf (northern Aegean Sea) (12), Gulf of Izmir (2)and GökovaGulf (southern Aegean Sea) (13) and off the coastof western Turkey by Meriç (1996) and Meriç et al. (2001b,2005, 2008b) (Fig. 1). Quite diversified benthic foraminiferalassemblages including 62 species were recognised at twolocalities, the Korthi and Kastro Gulfs (Andros Island), byTriantaphyllou et al. (2005) and Koukousioura et al. (2011)(14). A total of 67 benthic foraminiferal species were docu-mented in the Gulf of Izmir (western Turkey, eastern AegeanSea) by Bergin et al. (2006) (2). The benthic foraminiferaldiversity of Gulf of Izmir was also studied by Meriç et al.(2012a) and Barut et al. (2013). A very detailed investigationin the Gulf of Kalloni (Lesvos Island, Greece) (15) has beencarried out by Debenay et al. (2005) who documented thepresence of 147 benthic foraminiferal species. Mineralogicaland geochemical features of coloured benthic foraminiferalwere investigated in different Aegean Sea sites including theGökçeada region (3), southwestern Bozcaada (4, 11), andeastern Mitilini (16) by Yalçın et al. (2008). A well-diversified benthic foraminiferal assemblage was reported inPamucak Cove (17) (NW Kusadasi) that was defined as anoasis for alien species in the Aegean Sea (Meriç et al. 2008c,2009a, 2010a) (Fig. 1). Accordingly, these authors foundsome Indo-Pacific taxa such as Pyramidulina catesbyi,Brizalina simpsoni, and A. lessonii and a total of 103 speciesthat belong to 59 genera. Eighty-seven benthic foraminiferal

species were recorded in the continental shelf of the BüyükMenderes River delta (western Turkey) by Avşar et al. (2009)(18) (Fig. 1). On the basis of 26 sediment samples collectedalong Dardanelles, Meriç et al. (2009b) (19) documented avery rich foraminiferal assemblages composed of 119 speciesbelonging to 74 genera. The presence and the life cycle of thesymbiont-bearing foraminifera Amphistegina lobifera werestudied in other parts of the central and northern Aegean Sealike Falasarna (20), Porto Heli (21), Mavro Lithari (22),Vravron (23), Kalamos (24), Andros Island (14), Katerini(25) N. Kallikratia (26) and Avdira (27) by Triantaphyllouet al. (2009, 2012) (Fig. 1). A total of 93 benthic foraminiferalspecies belonging to 45 genera have been identified in thecoastal sites of Alykes Kitros (28), Lafrouda Thrace (27) andVravron Attica (23) in the western part of the Aegean Seathrough analysis of boreholes (Koukousioura et al. 2012).Bassler-Veit et al. (2013) identified 29 species of benthicforaminifera in salt pans, hypersaline lagoons, and salt lakesof the Gulf of Saros (6, 10, 29, 30, 31), the Biga Peninsula (32)and Gokçeada Island (3). The occurrence of benthic forami-niferal taxa in the eastern part of the Aegean Sea has been welldocumented by Meriç et al. (2004b) (Fig. 1), who recognised244 benthic foraminiferal species belonging to 124 genera. Inthis paper, several eastern Aegean Sea areas have been sur-veyed including: Saros Bay (6, 10, 29–31), around Gokçeadaisland (3), around Bozcaada (4, 11), Canakkale (5, 33), theGulf of Edremit (7, 12, 34), Dikili Bay (35), Candarli Bay(36), Karaburun Peninsula (37), the Gulf of Kusadasi (38),Gulluk Bay (39), the Gulf of Gökova (13), the Gulf of Datca(40) and Marmaris Bay (41) (Fig. 1). Further observations ofabnormal specimens of P. pertusus and P. planatus weredocumented in Illica Bay (42) by Meriç et al. (2012b)(Fig. 1). The same authors documented the occurrence of80 benthic foraminiferal species belonging to 45 genera. Inan environmental investigation carried out in the Camaltisaltpan (43), Meriç et al. (2010b) reported a total of 13species belonging to 8 genera. The benthic foraminiferalassemblages in the north coast of the Karaburun peninsula(37, 42) reveal the occurrence of 84 species (Meriç et al.2012c) (Fig. 1). In the same area, the alien foraminiferalspecies Polymorphina fistulosa was found by Meriç et al.(2012d). Several central-western Aegean Sea localities in-cluding Falasarna, Chrissi, Porto Heli, Mavro Lithari,Glyfada, Varkiza, Faliro, Perama, Vravron, Kalamos, theAndros-Korthi Gulf, Kastro Gulf and N. Kallikratia werestudied for documenting the distributional ranges of sixalien species (A. lobifera, S. orbicularis, C. hemprichii,Triloculina fichteliana, Planogypsina acervalis, andCymbaloporetta plana) (Koukousioura et al. 2010). Morerecently, a total of 96 agglutinated foraminiferal speciesbelonging to 51 genera were recognised along a bathymet-ric transect in Saros Bay (29, 31) (Frontalini et al. 2014)(Fig. 1).

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Materials and methods

A total of 18 samples were collected by Shipeck grab on boardthe R/V “Koca Piri Reis” during March 2003 along an innerneritic-middle bathyal transect (15–500 m) in Saros Bay(Fig. 2; Appendix 1). Once on board, surficial sediments (0–2 cm) were immediately washed through a 63-μm sieve, andpreserved in ethanol with Rose Bengal (2 g/L). Samples weresubsequently washed again, dried, and split into aliquots usinga modified Otto microsplitter in laboratory. At least 300 spec-imens of calcareous and of agglutinated living and deadforaminifera (total assemblages) were, when feasible, pickedfrom each sample (fraction >125 μm) and taxonomicallyidentified largely following both the generic classificationsof Loeblich and Tappan (1987) and the specificclassification of Cimerman and Langer (1991), Sgarrella andMoncharmont Zei (1993), Jones (1994), Rasmussen (2005),AGIP (1982), Parker (1958) and Jorissen (1988). The totalassemblages that comprise both living and dead foraminiferamight include autochthonous taxa as well as allochthonousones that might have been transported from other environ-ments. Broken specimens were identified to the genus level (ifpossible) and measured under each genus as unknown secies.If the genus was not recognisable, these specimens were nottaken into account. For the systematic classification schemeproposed for the benthic foraminiferal genera and thesuprageneric taxonomic categories adopted in this study, wehave substantially followed the suprageneric classificationsuggested by Hayward et al. (2013). On the basis of thisclassification, 17 orders are defined (Allogromiida,Astrorhizida, Lituolida, Loftusiida, Textulariida,Trochamminina, Miliolida, Robertinida, Buliminida,

Carterinida, Fusulinida, Globigerinida, Involutinida,Lagenida, Rotaliida, Silicoloculinida, and Spirillinida). TheFusulinida and Involutinida are two extinct orders.

Results and discussion

The Saros Bay area shows a very highly diversified benthicforaminiferal fauna, accordingly a total of 267 species weretaxonomically identified along a neritic to mid-bathyal tran-sect in water depths ranging from 15 to 500 m (Figs. 1, 2).These figures outnumber the diversity values documented inprevious investigations in the same area (Meriç et al. 2002b,2004a, 2005; Bassler-Veit et al. 2013). Benthic foraminiferalspecies/genera vary along a depth transect and also alreadyshown by Phleger (1960) in the Gulf of Mexico, it is possibleto define benthic foraminiferal assemblages typical for differ-ent bathymetric ranges. This has been partly already carried

Fig. 3 1 Ammodiscus gullmarensis Höglund, 1948. 2 Usbekistaniacharoides (Jones & Parker, 1860). 3 Adercotryma glomerata (Brady,1878). 4 Lagenammina atlantica (Cushman, 1944). 5, 6 Saccorhizaramosa (Brady, 1879). 7 Technitella legumen Norman, 1878. 8Ammolagena clavata (Jones & Parker, 1860). Scale bar 100 μm

Fig. 2 Geographical locations of the samples collected along the bathy-metric transect in Saros Bay. Isobaths are plotted at 200-m intervals

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out on the agglutinated foraminiferal assemblages byFrontalini et al. (2014) with the same samples that are herecoupled with the calcareous foraminiferal counterpart. In par-ticular, different bathymetric assemblages occurring at a cer-tain water depth can be recognised, such as the inner-middleneritic assemblages (ca. <100 m depth), outer neritic assem-blages (ca. 100–200 m), and upper bathyal assemblages (200–500m). Themost abundant taxa were pictured (Figs. 3, 4, 5, 6,7, 8, 9, and 10). It must be underlined that this study hasfocused on the total assemblages (living and dead foraminif-era) and therefore it cannot be excluded that the occurrence ofallochthonous taxa might have been transported from otherenvironments.

The inner neritic assemblages are mainly represented byBigenerina nodosaria , Lagenammina at lant ica ,Lagenammina d i f f l ug i f o rm i s , Sahu l i a con i ca ,

Spiroplectammina sagittula, Spiroplectinella wrightii,Textularia agglutinans, Textularia pseudogramen, Ammoniaparkinsoniana, Ammonia spp., Asterigerinata mamilla,Asterigerinata spp., Lobatula lobatula, Cibicides refulgens,Discorbinella bertheloti, Neoconorbina terquemi,Neoconorbina posidonicola, Planorbulina mediterranensis,Planorbulina variabilis, Planorbulina sp. nov., Buccellagranulata, Ammonia tepida, Cibicides pachydermaGlobocassidulina spp., Miliolinella subrotunda, Miliolinellaspp., Quinqueloculina spp., Sigmoilina costata, Rosalinabradyi, Rosalina floridensis, Rosalina globularis, Rosalinaobtusa, Rosalina spp., Rosalina vilardeboana, Cassidulinacarinata, Cassidulina laevigata, Cassidulina obtusa,Cassidulina spp., Elphidium spp., Elphidium complanatum,Astrononion stelligerum, Melonis barleeanus, and Reussellaspinulosa (Appendices 2, 3).

Fig. 4 1 Bigenerina nodosaria d’Orbigny, 1826. 2 Reophaxdentaliniformis Brady, 1881. 3 Textularia pala Czjzek, 1848. 4Spiroplectammina sagittula (Defrance, 1824). 5 Siphotextularia concava(Karrer, 1868). 6 Veleroninoides scitulus (Brady, 1881). 7Ammoglobigerina globigeriniformis (Parker & Jones, 1865). 8Recurvoidella bradyi (Robertson, 1891). 9 Trochammina nana (Brady,1881). Scale bar 100 μm.

Fig. 5 1, 2 Cycloforina contorta (d’Orbigny, 1846). 3 Pseudotriloculinalaevigata (d’Orbigny, 1826). 4, 5 Adelosina colomii (Le Calvez & LeCalvez, 1958). 6, 7 Triloculina gibba d’Orbigny, 1825. 8a, bSpiroloculina depressa d’Orbigny, 1826. 9 Triloculina tricarinatad’Orbigny, 1826. 10 Triloculina tricarinata d’Orbigny, 1826. 11a, bQuinqueloculina parvula Schlumberger, 1894. 12a, b Pyrgoella sphaera(d’Orbigny, 1839). 13, 14 Miliolinella subrotunda (Montagu, 1803).Scale bar 100 μm

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The outer neritic assemblages are instead characterised byBigenerina nodosaria, Eggerelloides scaber, Lagenamminaatlantica, Psammosiphonella spp., Pseudoclavulina crustata,Reophax scorpiurus, Usbekistania charoides, Rhabdamminaabyssorum, Saccorhiza ramosa, Siphotextularia concava,Spiroplectammina sagittula, Textularia agglutinans,Textularia pseudogramen, Lobatula lobatula, Cibicidesrefulgens, Planorbulina variabilis, Neoconorbinapo s i d on i c o l a , G l o bo ca s s i d u l i n a s ubg l o bo s a ,Globocassidulina spp., Quinqueloculina spp., Hyalineabalthica, Rosalina globularis, Cassidulina laevigata,Cassidulina obtusa, Astrononion stelligerum, Melonisbarleeanus, Sphaeroidina bulloides, Bulimina marginata,Bulimina mexicana, Uvigerina mediterranea, Buliminaaculeata, Bulimina gibba, Bolivina cf. dilatata, Bolivina

dilatata, Bolivina spp. Brizalina alata, Bolivina spathulata(Appendices 2, 3).

The deepest assemblages, namely upper bathyal assem-blages, are mainly dominated by Bigenerina nodosaria,Cribrostomoides subglobosum, Eggerelloides scaber,Martinottiella communis, Psammosiphonella spp. ,Pseudoclavulina crustata, Reophax fusiformis, Reophaxscorpiurus, Usbekistania charoides, Rhabdamminaabyssorum, Saccammina sphaerica, Saccorhiza ramosa,Spiroplectammina sagittula, Lobatula lobatula, Discorbinellabertheloti, Miliolinella subrotunda, Hyalinea balthica,Cassidulina carinata, Cassidulina laevigata, Elphidiumcomplanatum, Astrononion stelligerum, Melonis barleeanus,Sphaeroidina bulloides, Bulimina marginata, Buliminamexicana, Uvigerina mediterranea, Bulimina aculeata,Chilostomella mediterranensis, Bolivina cf. dilatata, Bolivinadilatata, Bolivina spp., Bolivina spathulata and Brizalinaalata (Appendices 2, 3).

On the basis of our data coupled with those available, weidentified 12 different orders of benthic foraminifera amongst

Fig. 6 1 Uvigerina mediterranea Hofker, 1932. 2 Reussella spinulosa(Reuss, 1850). 3, 4 Bulimina marginata d’Orbigny, 1826. 5a, b Buliminastriata d’Orbigny, in Guérin-Méneville, 1843. 6, 7 Bulimina aculeatad’Orbigny, 1826. 8, 9 Bulimina elongata d’Orbigny, 1846. 10 Buliminagibba Fornasini, 1902. 11, 12 Fursenkoina complanata Egger, 1893. 13Bolivina striatula Cushman, 1922. 14 Bolivina cf. dilatata Reuss, 1850.15 Bolivina dilatataReuss, 1850. 16a, b Bolivina artaMacFadyen, 1930.17 Globobulimina auriculata (Bailey, 1851). 18, 19 Globobuliminapseudospinescens (Emiliani 1949). 20, 21 Brizalina spathulata(Williamson, 1858). Scale bar 100 μm

Fig. 7 1, 2 Elphidium complanatum (d’Orbigny, 1839). 3a, b Elphidiumaculeatum (d’Orbigny, 1846). 4, 5 Melonis barleeanus (Williamson,1858). 6a, b)Elphidium granosum (d’Orbigny, 1826). 7 Cassidulinacarinata Silvestri, 1896. 8, 9a, b Gyroidina orbicularis d’Orbigny,1826. 10, 11 Rosalina bradyi Cushman, 1915. 12, 13 Hyalinea balthica(Schröter, 1783). Scale bar 100 μm

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the 14 living ones as suggested by Hayward et al. (2013) aswell as the planktonic order Globigerina in the Aegean Sea.The drying of the samples does not allow the preservation ofthe informally defined naked-foraminifera and therefore, theorder Allogromiida is not considered here. We recognise theAstrorhizida, Lituolida, Loftusiida, Textulariida,Trochamminina (which includes the Carterinida in the classi-fication of Kaminski (2004)), Miliolida, Robertinida,Spirillinida, Lagenida, Buliminida, and Rotaliida (Table 1;Appendix 4).

Three superfamilies (Astrorhizoidea Saccamminoidea andHippocrepinoidea), eight families (Bathysiphonidae,Rhabdamminidae , Saccaminidae , Ast rorhiz idae ,P s ammo s p h a e r i d a e , H em i s p h a e r amm i n i d a e ,Hippocrepinidae, Cri thioninidae) and 19 genera(Bathysiphon, Psammosiphonella, Rhabdammina,Marsipella, Lagenammina, Saccammina, Technitella,Thurammina, Tholosina, Iridia, Webbinella, Bigenerina,Psammosphaera, Hemisphaerammina, Crithionina,Botellina, Hyperammina, Saccorhiza and Jaculella) havebeen identified within the Order Astrorhizida (Table 1).The Order Lituolida is represented by ten superfamilies

Fig. 9 1a, b Cibicides dutemplei (d’Orbigny, 1846). 2, 3 Cibicidespachyderma (Rzehak, 1886). 4–6 Buccella granulata (Di Napoli Alliata,1952). 7–9 Asterigerinata mamilla (Williamson,1858). 10, 11 Planorbulinavariabilis (d’Orbigny, 1826). 12, 13 Discorbinella bertheloti (d’Orbigny,1839). 14–16 Neoconorbina terquemi (Rzehak, 1888). Scale bar 100 μm

Fig. 10 1a, b Planorbulina sp. nov. 2 Planorbulina sp. nov. Scale bar100 μm

Fig. 8 1a, b Elphidium macellum (Fichtel & Moll, 1798). 2 Ammoniabeccarii (Linnaeus, 1758).. 3, 4a, b Ammonia cf. parkinsoniana(d’Orbigny, 1839). 5, 6a, b Ammonia cf. tepida (Cushman, 1926). Scalebar 100 μm

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Table 1 Benthic foraminiferal taxa (order, superfamily, family, and genus) and species number recognized in the Aegean Sea

Order Superfamily Family Genus Species

Astrorhizida Astrorhizoidea Bathysiphonidae Bathysiphon 1

Psammosiphonella 1

Rhabdamminidae Rhabdammina 4

Marsipella 2

Saccaminidae Lagenammina 3

Saccammina 1

Technitella 1

Thurammina 2

Tholosina 1

Iridia 1

Webbinella 1

Astrorhizidae Bigenerina 3

Psammosphaeridae Psammosphaera 1

Hemisphaeramminidae Hemisphaerammina 1

Saccamminoidea Crithioninidae Crithionina 1

Hippocrepinoidea Hippocrepinidae Botellina 1

Hyperammina 4

Saccorhiza 1

Jaculella 1

Lituolida Ammodiscoidea Ammodiscidae Ammodiscus 7

Glomospira 2

Usbekistania 1

Hormosinelloidea Ammolagenidae Ammolagena 1

Hormosinoidea Hormosinidae Reophax 9

Cuneata 1

Hormosina 1

Reophacidae Leptohalysis 1

Haplophragmioidea Discamminidae Ammoscalaria 3

Discammina 1

Glaphyrammina 1

Haplophragmoididae Haplophragmoides 2

Evolutinella 1

Veleroninoides 2

Ammosphaeroidinidae Recurvoides 1

Adercotryma 1

Ammosphaeroidina 1

Lituoloidea Lituolidae Cribrostomoides 4

Ammobaculites 1

Placopsilinidae Placopsilina 1

Lituotuboidea Lituotubidae Lituotuba 1

Spiroplectamminoidea Spiroplectamminidae Spiroplectinella 3

Spiroplectammina 1

Nouriidae Nouria 1

Verneuilinoidea Verneuilinidae Connemarella 1

Dorothia 3

Gaudryina 3

Verneuilinulla 2

Loftusioidea Cyclamminidae Cyclammina 1

Rzehakinoidea Rzehakinidae Miliammina 1

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Table 1 (continued)

Order Superfamily Family Genus Species

Loftusiida Coscinophragmatoidea Haddoniidae Haddonia 1

Textulariida Textularioidea Pseudogaudryinidae Pseudoclavulina 1

Textulariidae Siphotextularia 7

Textularia 16

Sahulia 1

Planctostoma 1

Eggerelloidea Eggerellidae Eggerelloides 1

Martinottiella 1

Karreriella 1

Eggerella 1

Valvulinidae Cribrogoesella 1

Cylindroclavulina 1

Clavulina 2

Trochamminina Trochamminoidea Trochamminidae Trochammina 3

Ammoglobigerina 1

Trochamminopsis 1

Polystomammina 1

Recurvoidella 1

Tritaxis 1

Lepidodeuterammina 1

Paratrochammina 1

Carterinida Carterinidae Carterina 1

Miliolida Cornuspiroidea Cornuspiridae Cornuspira 4

Nubecularioidea Ophthalmidiidae Spirophthalmidium 2

Ophthalmidium 2

Cornuloculina 1

Edentostomina 2

Nubeculariidae Nubeculina 1

Nubecularia 1

Fischerinidae Vertebralina 2

Nodophthalmidium 2

Wiesnerella 1

Trisegmentina 1

Planispirina 1

Milioloidea Cribrolinoididae Adelosina 17

Hauerinidae Biloculinella 4

Schlumbergerina 1

Cycloforina 5

Hauerina 1

Miliolinella 10

Massilina 3

Lachlanella 2

Pseudolachlanella 1

Pseudotriloculina 4

Pyrgo 13

Pyrgoella 3

Quinqueloculina 42

Triloculina 15

Triloculinella 1

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Table 1 (continued)

Order Superfamily Family Genus Species

Sigmoilina 5

Sigmoilopsis 2

Spirosigmoilina 2

Siphonaperta 5

Affinetrina 1

Articulina 6

Parrina 1

Cruciloculina 1

Sigmoihauerina 1

Nummoloculina 1

Pseudoschlumbergerina 1

Wellmanellinella 1

Spiroloculinidae Spiroloculina 20

Nummulopyrgo 1

Soritoidea Peneroplidae Peneroplis 3

Coscinospira 2

Euthymonacha 1

Monalysidium 1

Soritidae Sorites 4

Amphisorus 1

Robertinida Robertinoidea Robertinidae Robertina 2

Ceratobuliminoidea Epistominidae Epistomina 1

Ceratobuliminidae Lamarckina 1

Rubratella 1

Ceratocancris 1

Spirillinida Spirillinidae Spirillinidae Spirillina 6

Patellinidae Patellina 1

Mychostomina 2

Lagenida Nodosarioidea Vaginulinidae Amphicoryna 8

Hemirobulina 1

Lenticulina 7

Neolenticulina 2

Saracenia 1

Astacolus 3

Marginulinopsis 3

Marginulina 3

Vaginulina 2

Vaginulinopsis 1

Nodosariidae Dentalina 8

Grigelis 2

Lingulina 5

Laevidentalina 6

Pyramidulina 4

Nodosaria 3

Frondicularia 2

Lagenidae Lagena 21

Pygmaeoseistron 1

Procerolagena 2

Reussoolina 1

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Table 1 (continued)

Order Superfamily Family Genus Species

Cerebrina 1

Cristellaria 1

Polymorphinoidea Polymorphinidae Laryngosigma 2

Globulina 5

Vasiglobulina 1

Polymorphina 4

Guttulina 1

Pseudopolymorphina 1

Sigmomorphina 1

Pyrulina 1

Ellipsolagenidae Favulina 4

Fissurina 19

Oolina 6

Parafissurina 10

Homalohedra 1

Lagenosolenia 5

Lagnea 1

Cushmanina 1

Palliolatella 2

Glandulinidae Glandulina 2

Buliminida Buliminoidea Buliminidae Bulimina 11

Globobulimina 5

Stainforthia 2

Trimosinidae Mimosina 1

Buliminellidae Buliminella 1

Uvigerinidae Trifarina 3

Uvigerina 11

Reussella 2

Siphogenerinoididae Sagrina 1

Rectuvigerina 1

Hopkinsina 1

Siphogenerina 1

Loxostomina 1

Spiroloxostoma 1

Delosinoidea Delosinidae Delosina 1

Fursenkoinoidea Fursenkoinidae Fursenkoina 5

Sigmavirgulina 2

Virgulopsis 1

Coryphostoma 2

Virgulina 1

Rutherfordoides 1

Bolivinoidea Bolivinidae Bolivina 23

Bolivinellina 1

Brizalina 3

Bolivinitoidea Bolivinitidae Bolivinita 1

Tortoplectellidae Tortoplectella 1

Cassidulinoidea Cassidulinidae Cassidulina 4

Cassidulinoides 1

Globocassidulina 4

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Table 1 (continued)

Order Superfamily Family Genus Species

Ehrenbergina 1

Rotaliida Discorbinelloidea Planulinoididae Discorbinella 1

Parrelloididae Cibicidoides 2

Pseudoparrellidae Eilohedra 1

Epistominella 1

Discorboidea Sphaeroidinidae Sphaeroidina 1

Discorbidae Buccella 2

Rotorbinella 1

Gavelinopsis 1

Strebloides 1

Tretomphalus 2

Discorbina 3

Discorbis 5

Rotorbis 1

Disconorbis 1

Crouchina 1

Orbitina 1

Bagginidae Valvulineria 4

Cancris 3

Baggina 1

Rosalinidae Neoconorbina 3

Rosalina 12

Planodiscorbis 1

Tretomphaloides 1

Mississippinidae Stomatorbina 2

Eponididae Eponides 3

Planopulvinulina 1

Neoeponides 1

Poroeponides 1

Chilostomelloidea Gavelinellidae Gyroidina 6

Anomalinoides 2

Hanzawaia 3

Chilostomellidae Chilostomella 2

Heterolepidae Heterolepa 1

Trichohyalidae Aubignyna 2

Alabaminidae Svratkina 1

Oridorsalidae Oridorsalis 1

Asterigerinoidea Asterigerinatidae Asterigerinata 4

Pninaella 1

Epistomariidae Nuttallides 1

Amphisteginidae Amphistegina 2

Planorbulinoidea Cibicididae Cibicides 9

Lobatula 1

Cyclocibicides 2

Planulinidae Hyalinea 1

Planulina 1

Planorbulinidae Planorbulina 3

Planogypsina 2

Caribeanella 1

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(Ammodiscoidea, Hormosinelloidea, Hormosinoidea,Haplophragmioidea, Li tuoloidea, Li tuotuboidea,Spiroplectamminoidea, Verneuilinoidea Loftusioidea andRzehakinoidea), 12 families (Ammodiscidae, Ammolagenidae,Ho rmos i n i d a e , Reophac i d a e , D i s c ammin i d a e ,Haplophragmoididae, Ammosphaeroidinidae, Lituolidae,Placopsilinidae, Lituotubidae, Spiroplectamminidae, Nouriidae,Verneuilinidae, Cyclamminidae and Rzehakinidae) and 30 gen-era (Ammodiscus, Glomospira, Usbekistania, Ammolagena,Reophax, Cuneata, Hormosina, Leptohalysis, Ammoscalaria,Discammina, Glaphyrammina, Haplophragmoides,Evolutinella, Veleroninoides, Recurvoides, Adercotryma,Ammosphaeroidina, Cribrostomoides, Ammobaculites,Placopsilina, Lituotuba, Spiroplectinella, Spiroplectammina,Nouria, Connemarella, Dorothia, Gaudryina, Verneuilinulla,Cyclammina and Miliammina) (Table 1).

The order Loftusiida is only represented by thesuperfamiliy Coscinophragmatoidea and the familyHaddoniidae and the genus Haddonia (Table 1).

The order Textulariida includes only two superfamilies(Textularioidea and Eggerelloidea), four families(Pseudogaudryinidae, Textulariidae, Eggerellidae andValvul in idae) and 12 genera (Pseudoclavul ina,Siphotextularia, Textularia, Sahulia, Planctostoma,Eggerelloides, Martinottiella, Karreriella, Eggerella,Cribrogoesella, Cylindroclavulina and Clavulina), whereasthe order Trochamminida contains one superfamily(Trochamminoidea), one family (Trochamminidae) andeight genera (Ammoglobigerina , Trochammina,Trochamminopsis, Polystomammina, Recurvoidella,Tritaxis Lepidodeuterammina, and Paratrochammina)(Table 1).

Table 1 (continued)

Order Superfamily Family Genus Species

Victoriellidae Carpenteria 1

Cymbaloporidae Cymbaloporetta 4

Nonionoidea Nonionidae Pullenia 4

Melonis 4

Nonionella 5

Astrononion 4

Haynesina 4

Nonion 4

Nonionoides 1

Rotalioidea Elphidiidae Elphidium 26

Cribroelphidium 3

Porosononion 1

Parrellina 1

Rotaliidae Ammonia 8

Challengerella 1

Pararotalia 1

Siphoninoidea Siphoninidae Siphonina 2

Glabratelloidea Glabratellidae Pileolina 1

Conorbella 4

Glabratellina 1

Planoglabratella 1

Schackoinella 1

Glabratella 1

Heronalleniidae Heronallenia 1

Acervulinoidea Acervulinidae Acervulina 1

Gypsina 1

Sphaerogypsina 1

Homotrematidae Miniacina 2

Nummulitoidea Nummulitidae Heterocyclina 1

Heterostegina 1

Incertae sedis Pulvinulina 1

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The order Carterinida is included within theTrochamminida in the classification of Kaminski (2004) andis represented by only one family, Carterinidae, and one genusCarterina (Table 1).

The order Miliolida encompasses 4 superfamilies(Cornuspiroidea, Nubecularioidea, Milioloidea andSoritoidea), 9 families (Cornuspiridae, Ophthalmidiidae,Nubeculariidae, Fischerinidae, Cribrolinoididae, Hauerinidae,Spiroloculinidae, Peneroplidae and Soritidae) and 47 genera(Cornuspira, Spirophthalmidium, Ophthalmidium,Cornuloculina, Edentostomina, Nubeculina, Nubecularia,Ver tebra l ina , Nodoph tha lmid ium, Wiesnere l la ,Trisegmentina, Planispirina, Adelosina, Biloculinella,Schlumbergerina, Cycloforina, Hauerina, Miliolinella,Ma s s i l i n a , L a c h l a n e l l a , P s e u d o l a c h l a n e l l a ,Pseudotriloculina, Pyrgo, Pyrgoella, Quinqueloculina,Triloculina, Triloculinella, Sigmoilina, Sigmoilopsis,Spirosigmoilina, Siphonaperta, Affinetrina, Articulina,Parrina, Cruciloculina, Sigmoihauerina, Nummoloculina,Pseudoschlumbergerina, Wellmanellinella, Spiroloculina,N ummu l o p y r g o , P e n e r o p l i s , C o s c i n o s p i r a ,Euthymonacha, Monalysidium, Sorites, and Amphisorus)(Table 1).

The orderRobertinida is represented by two superfamilies(Robertinoidea and Ceratobuliminoidea ), three families(Robertinidae, Epistominidae, and Ceratobuliminidae) andfive genera (Robertina, Epistomina, Lamarckina, Rubratella,and Ceratocancris).

The order Spirillinida is composed of one superfamily(Spirillinidae), two families (Spirillinidae and Patellinidae)and three genera (Spirillina, Patellina and Mychostomina)(Table 1).

Tw o s u p e r f a m i l i e s (N o d o s a r i o i d e a a n dPolymorphinoidea) , six famil ies (Vaginulinidae,Nodo s a r i i d a e , Lag en i d a e , P o l ymo r p h i n i d a e ,Ellipsolagenidae and Glandulinidae) and 41 genera(Amphicoryna, Hemirobulina, Lenticulina, Neolenticulina,Saracenia, Astacolus, Marginulinopsis, Marginulina,Vaginulina, Vaginulinopsis, Dentalina, Grigelis, Lingulina,Laevidentalina, Pyramidulina, Nodosaria, Frondicularia,Lagena, Pygmaeoseistron, Procerolagena, Reussoolina,Cerebrina, Cristellaria, Laryngosigma, Globulina,Va s i g l o b u l i n a , P o l y m o r p h i n a , G u t t u l i n a ,Pseudopolymorphina, Sigmomorphina, Pyrulina, Favulina,Fissurina, Oolina, Parafissurina, Homalohedra,Lagenosolenia, Lagnea, Cushmanina, Palliolatella andGlandulina) are recognised within the order Lagenida(Table 1).

On the other hand, the order Buliminida contains sixs u p e r f am i l i e s (B u l im i n o i d e a , D e l o s i n o i d e a ,Fursenkoinoidea, Bolivinoidea, Bolivinidae,andCassidulinoidea), 11 families (Buliminidae, Trimosinidae,Buliminellidae, Uvigerinidae, Siphogenerinoididae,

Delosinidae, Fursenkoinidae, Bolivinidae, Bolivinitidae,Tortoplectellidae and Cassidulinidae) and 30 genera(Bulimina, Globobulimina, Stainforthia, Mimosina,Buliminella, Trifarina, Uvigerina, Reussella, Sagrina,Rectuvigerina, Hopkinsina, Siphogenerina, Loxostomina,Spiroloxostoma, Delosina, Fursenkoina, Sigmavirgulina,Virgulopsis, Coryphostoma, Virgulina, Rutherfordoides,Bo l i v ina , Bo l i v ine l l ina , Br i za l ina , Bo l i v in i ta ,Tor top l e c t e l l a , Cas s i du l i na , Cas s i du l i no ide s ,Globocassidulina, and Ehrenbergina) (Table 1).

The order Rotaliida is the most diversified taxon and ismade up of 11 superfamilies (Discorbinelloidea,Discorboidea, Chilostomelloidea, Asterigerinoidea,Planorbulinoidea, Nonionoidea, Rotalioidea, Siphoninoidea,Glabratelloidea, Acervulinoidea and Nummulitoidea), 32families (Planulinoididae, Parrelloididae, Pseudoparrellidae,Sphaeroidinidae, Discorbidae, Bagginidae, Rosalinidae,Mis s i s s ipp in idae , Epon id idae , Gave l ine l l i dae ,Chilostomellidae, Heterolepidae, Trichohyalidae,Alabaminidae, Oridorsal idae, Aster iger inat idae,Epistomariidae, Amphisteginidae, Cibicididae, Planulinidae,Planorbulinidae, Victoriell idae, Cymbaloporidae,Nonionidae, Elphidiidae, Rotaliidae, Siphoninidae,Glabra te l l idae , Heronal leni idae , Acervul in idae ,Homotrematidae, and Nummulitidae) and 78 genera(Discorbinella, Cibicidoides, Eilohedra, Epistominella,Sphaeroidina, Buccella, Rotorbinella, Gavelinopsis,Strebloides, Tretomphalus, Discorbina, Discorbis, Rotorbis,Disconorbis, Crouchina, Orbitina, Valvulineria, Cancris,Baggina, Neoconorbina, Rosalina, Planodiscorbis,Tretomphaloides, Stomatorbina, Eponides, Planopulvinulina,Neoeponides, Poroeponides, Gyroidina, Anomalinoides,Hanzawaia, Chilostomella, Heterolepa, Aubignyna,Svratkina, Oridorsalis, Asterigerinata, Pninaella, Nuttallides,Amphistegina, Cibicides, Lobatula, Cyclocibicides, Hyalinea,Planulina, Planorbulina, Planogypsina, Caribeanella,Carpenteria, Cymbaloporetta, Pullenia, Melonis, Nonionella,Astrononion, Haynesina, Nonion, Nonionoides, Elphidium,Cribroelphidium, Porosononion, Parrellina, Ammonia,Challengerella, Pararotalia, Siphonina, Pileolina,Conorbella, Glabratellina, Planoglabratella, Schackoinella,Glabratella, Heronallenia, Acervulina, Gypsina,Sphaerogypsina, Miniacina, Heterocycl ina, andHeterostegina) (Table 1).

On the basis of the available foraminiferal studies pub-lished on the Aegean Sea coupled with our data from SarosBay, a very high number of foraminiferal species is found tooccur in this Mediterranean basin (Appendix 4). In particular,a total of 799 species belonging to 276 genera, 93 families and43 superfamilies have been recorded in the Aegean Sea. Itmust be, however, underlined that most of the consideredstudies were based on the total assemblages and thereforemight have included allochthonous taxa from other

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environments and the high diversity might most probablyrepresent a time-averaged biodiversity hotspot.

These values are remarkably higher than those documentedin previous foraminiferal papers carried out in different partsof the Mediterranean Sea. Accordingly, a total of 288 moderncarbonate shelf benthic foraminifera were recognised byMilk-er and Schmiedl (2012) in three areas of the Western Medi-terranean Sea (Alboran Platform, Oran Bight and the south-west shelf of Mallorca), whereas 334 species weredocumented by Cimerman and Langer (1991) for theAdriatic and Tyrrhenian Seas. An unexpected result of ourcompilation is the fact that the foraminiferal diversity of theAegean Sea is comparable to that of the highly diverse SahulShelf and Timor Sea, where Loeblich and Tappan (1994)reported 946 species. Although the determination of the totalforaminiferal diversity in the Mediterranean Sea remains adifficult task because the deep-water agglutinated and nakedforms are still largely unstudied, we estimate that the totalnumber of foraminiferal species might be well in excess of1,000.

Conclusions

Benthic foraminiferal species diversity has been investigatedalong a neritic to mid-bathyal transect (15–500 m) in SarosBay. Accordingly, a total of 267 species belonging to 115genera have been documented. Benthic foraminiferal taxavary along a depth transect and it is, therefore, possible todefine benthic foraminiferal assemblages typical for differentbathymetric ranges like the inner-middle neritic assemblages(ca. <100 m depth), outer neritic assemblages (ca. 100–200 m), and upper bathyal assemblages (200–500 m). Onthe basis of the available foraminiferal literature on the Aege-an Sea coupled with our data from Saros Bay, a very highnumber of foraminiferal species occur in this part of theMediterranean Sea. In particular, we estimate that a total of799 species belonging to 276 genera, 93 families and 43superfamilies have been recorded in the Aegean Sea. Thesevalues are remarkably higher than those documented in pre-vious foraminiferal studies carried out in other sectors of theMediterranean Sea. Although our estimate is a conservativeone, and the determination of the total foraminiferal diversityin the Mediterranean Sea remains an unfinished task, it mightbe speculated that this number might be well in excess of athousand.

Acknowledgments The research for this paper was partially madepossible by the financial support from the PRIN 2010–2011 Ministerodell’Istruzione, dell’Università e della Ricerca (MIUR) (protocollo2010RMTLYR). MAK thanks A.E. Aksu (Memorial University of New-foundland) and the Captain and crew of the R/V “Piri Reis” for theopportunity to collect samples in Saros Bay. The authors are also gratefulto Philippe Recourt for managing the SEM picture at University of Lille

1. We are particularly grateful to two anonymous reviewers for theirthoughtful and valuable comments that have greatly improved the paper.

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