Geologia CroaticaGeologia Croatica AB STRA CT This study represents a detailed micropalaeontological investigation of the composition and diversity of planktonic foraminiferal assemblages from the upper Turonian to Maastrichtian interval of two deep exploration wells (Al Mahr-1 and Palmyra-1) in the Palmyride area of Syria. In combination with lithostratigraphic analysis, this detailed biostrati- graphic study provided important new palaeoecological and palaeoclimatic interpretations and insights into the na- ture of deposition along the northern passive margin of Gondwana during the Late Cretaceous. The investigated strata belong to three lithostratigraphic units (from base to top): the upper part of the Judea Forma- tion (upper Turonian–lowermost Santonian), the Soukhne Formation (Santonian–lower Campanian), and the Shiran- ish Formation (upper Campanian–Maastrichtian). The results represent the fi rst detailed determination of planktonic foraminifera from the Palmyride region. The presence of rich and diverse foraminiferal associations enabled the es- tablishment of the following nine late Turonian to Maastrichtian biostratigraphic zones, based on documented index- taxa and/or the entire microfossil assemblages: I) Biozone I; II) Biozone II; III) Biozone III; IV) Contusotruncana plummerae Zone; V) Biozone V; VI) Globotruncanella havanensis Zone; VII) Pseudoguembelina palpebra Zone; VIII) Racemiguembelina fructicosa Zone; and IX) Abathomphalus mayaroensis Zone. The late Turonian to early Campanian foraminiferal assemblages (biozones I–IV) are dominated by opportunistic taxa (r-strategists) and sug- gest a generally fl uctuating subtropical climate and deposition an outer shelf environments. The well­preserved and highly diversifi ed late Campanian to Maastrichtian foraminiferal assemblages (biozones V–IX) imply the presence of a well­stratifi ed water column, tropical to subtropical climate, and deposition in outer shelf to upper bathyal envi­ ronments. A decrease in the number of globotruncanid species during the late Maastrichtian indicates a less stratifi ed water column and unfavourable environmental conditions for K-strategists. The common occurrence of phosphate grains in the Soukhne Formation (Santonian–lower Campanian) represents an important indicator of specifi c geological and palaeoenvironmental conditions, such as oxygen defi ciency, up­ welling and transgression. These conditions support the interpretation of the domination by opportunistic planktonic foraminiferal taxa (heterohelicids and muricohedbergellids) in Biozone II. Keywords: Planktonic foraminifera, Biostratigraphy, Phosphatic grains, Upper Turonian–Maastrichtian, Palmyride region, Syria Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria  Gabrijela Pecimotika1, Blanka Cvetko Tešović2 and Vlasta Premec Fućek1 1 E&P Research Laboratory Department, Field Engineering and Operations Sector, INA-Industrija nafte d.d., Exploration and Production BD, Lovinčićeva bb, 10 002 Zagreb, Croatia; (gabrijela.pecimotika@ina.hr; vlasta.premec­fucek@ina.hr) 2 Department of Geology, Faculty of Science, University of Zagreb, Horvatovac 102a, 10 000 Zagreb, Croatia; (bcvetko@geol.pmf.hr) doi: 10.4154/gc.2014.07 Geologia Croatica 67/2 87–110 10 Figs. Zagreb 2014 1. INTRODUCTION The Palmyride area is part of the northern Arabian platform (Fig. 1). The Arabian platform was located on the northern passive margin of Gondwana bordering the Tethys Ocean for most of the Phanerozoic. The Palmyride fold belt was established at the site of an inverted Mesozoic rift basin and Geologia Croatica 67/2Geologia Croatica 88 developed as a linear trough genetically related to the Le- vantine margin rift system, which formed along a Gondwana Proterozoic suture zone (BREW, 2001). Ongoing extension produced a 6 km thick and 200 km laterally extensive Pal- aeozioc and Mesozoic sedimentary succession (BREW, 2001). Tectonic evolution of the area has been strongly in- fluenced by geological activity along the Arabian plate boun­ daries: the Dead Sea transform fault to the west, the Bitlis suture and East Anatolian fault to the north, and the Zagros suture to the east (Fig. 1). This study focuses on the Upper Cretaceous (upper Tu- ronian–Maastrichtian) succession from two deep exploration wells Al Mahr-1 and Palmyra-1, and was aimed at age de- termination and correlation (litho- and biostratigraphic) of strata based on their microfossil assemblages (mainly plank- tonic and some benthic foraminifera). Planktonic foraminif- era have been abundant in most oceanic environments since their appearance in the Middle Jurassic and are the most commonly used microfossil group for biostratigraphic zona- tion and reconstruction of past sea surface-water conditions and palaeoclimate (HEMLEBEN et al., 1989; MURRAY, 1991). Many studies have focused on biostratigraphic evolution of Cretaceous planktonic foraminifera (e.g., ROBASZYN- SKI et al., 1984; 2000; CARON, 1985; SLITER, 1989; RO- BASZYNSKI & CARON, 1995; PETRIZZO, 2003; PRE­ MO LI SILVA & VERGA, 2004; SARI, 2006, 2009; BABA ZADEH et al., 2007; HUBER et al., 2008; PETRIZZO et al., 2011; GARDIN et al., 2012). Previous biostratigraphic investigations of the Upper Cretaceous deposits in the Palmy ride area were undertaken on material from deep ex- ploration wells by LUČIĆ (2001), STANKOVIĆ et al. (2003; 2005), and DACER et al. (2007), but not in a great detail. The main purpose of this paper is to establish planktonic foraminiferal zonation of the upper Turonian–Maastrichtian succession of the Palmyride area based on the microfossil assemblages and/or index taxa present in order to improve palaeoenvironmental interpretations of deposition in the Palmyride basin during the late Cretaceous. The biostrati- graphic zonation is compared with the regional Tethyan zo- Figure 1: Map showing major tectonic zones in Syria and the surrounding areas (modified from LITAK et al., 1998) and the study area in the Palmyride region. A circle represents the approximate location of the Hayan block with the Al Mahr-1 (1) and Palmyra-1 (2) exploration wells; distance between the wells is about 50 km. Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 89 nation. Documenting variations in planktonic foraminiferal assemblages and any associated lithological changes are crit- ical for making palaeoclimatic and palaeoceanographic in- terpretations. Santonian to early Campanian sediments rich in phosphate grains are examined here as an important indi- cator of specific geological and palaeoenvironmental condi- tions. The data obtained are compared to those from other coeval regional successions in order to establish the signifi- cance of the Palmyride strata, as part of the Arabian platform, for better understanding of the sedimentary evolution of the broader Tethyan region and its response to global environ- mental changes. 2. GEOLOGICAL SETTING Regionally the investigated area is also known as the Hayan exploration block (Fig. 1), which is located in the Palmyride area, an intracontinental transpressive mountain range (LUČIĆ, 2001). The Palmyrides represent the most distinct tectonic and structural unit in central Syria as a zone of sub- dued topography that extends from the Dead Sea Fault Zone to the west, and disappears to the east at the Euphrates Gra- ben or depression (Fig. 1). The Palmyrides are 400 km long and 100 km wide, stretching southwest–northeast across Syria with a maximum altitude of around 1300 m (LUČIĆ & FORŠEK, 2000; BREW, 2001; BREW et al., 2001; HER- NITZ KUČENJAK et al., 2006; WOOD, 2011). In the Palmyride area, Mesozoic deposits of Early Tri- assic to Late Cretaceous age were observed in all deep wells (LUČIĆ et al., 2002). Unlike the Upper Triassic and Jurassic deposits (maximum 700 m thick), which can either exhibit substantially reduced thickness (to 200 m minimum) or be absent in some places due to erosion or non-deposition, the Cretaceous strata are present throughout the region (approx. 800 m thick). The oldest deposits exposed on the surface are Upper Triassic evaporites interbedded with shales. Jurassic deposits are represented by different varieties of carbonate rocks, and Lower Cretaceous deposits consist of dolomites and limestones with rare interbeds of anhydrite and shale. In the Late Cretaceous there was a deepening of the deposi- tional system, which resulted in the deposition of shales and marly limestones with a gradual increase in the amount of marl up­section (PONIKAROV, 1966a, b; LUČIĆ et al., 200. For the purpose of the Syrian Petroleum Company (SPC), the investigated Upper Cretaceous succession is sub- divided into three lithostratigraphic units (Figs. 2, 3): 1) the upper part of the Judea Formation (upper Turonian–lower- most Santonian); 2) the Soukhne Formation (Santonian– lower Campanian); and 3) the Shiranish Formation (upper Campanian–Maastrichtian); (MOUTY & AL-MALEH, 1983). The Judea Formation is represented by limestones and dolomitic limestones with thin intercalations of yellow to brownish yellow marl. The Soukhne Formation is char- acterized by calcareous horizons in the lower part, and by clayey limestones, marls and phosphatic deposits in the up- per part. Argillaceous limestones, marls, chert and ovoid cal- careous concretions (10–30 cm in diameter) are present in the Shiranish Formation. 3. MATERIAL AND METHODS The foraminiferal study is based on analyses of 81 samples of Upper Cretaceous deposits obtained as drill cuttings from two deep exploration wells (Al Mahr-1 and Palmyra-1) drilled in the Hayan exploration block in the Palmyride area. Samples of drill cuttings from mud samples were collected every 5–10 metres. Most of the analyzed samples contain very well preserved planktonic and benthic foraminifera. Samples for micropalaeontological analyses were disag- gregated in tap water and diluted with hydrogen peroxide, then washed through 63 mm, 125 mm, and 160 mm sieves, dried and examined on an Olympus SZX16 stereomicro- scope. Representative aliquots of approximately 300 plank- tonic foraminiferal specimens were counted for quantitative planktonic foraminiferal analyses. The term “„dominant” was used for species that constitute more than 10% of the planktonic foraminiferal assemblage, whereas the terms “„common”, “„fewand “„rare” refer to species comprising 3–10%, 1–3%, and <1% of the assemblage, respectively. Plankton/benthic ratios were determined for each biozone on at least 300 specimens from the entire foraminiferal as- semblages in >63 mm grain fraction and were used for pal- aeoecologic and palaeoenvironmental interpretations. Petrographic thin­sections were made of 35 samples throughout the Upper Cretaceous interval for the purpose of lithological interpretation. Prepared petrographic thin-sec- tions were stained with Alizarin red – S after the method of EVAMY & SHEARMAN (1962) in order to distinguish car- bonate minerals. A detailed study of foraminiferal morphol- ogy was performed on a scanning electron microscope (SEM). The overall preservation of foraminifera is good al- though their original calcite shells have been recrystallized. The micropalaeontological investigation was focused on the vertical distribution, diversity and composition of mainly planktonic and less abundant benthic foraminiferal assem- blages (Figs. 2–3) according to GAWOR­BIEDOWA (1984), LOEBLICH & TAPPAN (1988), ISMAIL (1992), BOLLI et al. (1994), LY & KUHNT (1994), KAIHO (1998), PREMOLI SILVA &VERGA (2004), ISMAIL et al. (2007), HAMPTON et al. (2007), and GASIŃSKI & UCHMAN (2009). The pres- ence of rich and diverse foraminiferal associations enabled biostratigraphic zonation of the Upper Turonian to Maas- trichtian deposits based on documented index-taxa and/or the entire microfossil assemblages. Standard planktonic fo- raminiferal zonation after ROBASZYNSKI et al. (1984), CARON (1985), ROBASZYNSKI & CARON (1995), PRE­ MOLI SILVA & SLITER (1994), ROBASZYNSKI et al. (2000), PREMOLI SILVA &VERGA (2004), SARI (2006, 2009), HUBER et al. (2008), OGG et al. (2008), PETRIZZO et al. (2011) AND PÉREZ­RODRÍGUEZ et al. (2012) have been used. A zonal sheme which emphasizes the lowest (LO) and the highest occurrences (HO) of index taxa and/or se- lected species in microfossil associations has been applied. Phosphatic grains of the Soukhne Formation deposits were also analyzed using SEM back-scattered electron im- aging (BSE) and energy dispersive X-ray analysis (EDX). The semi-quantitative X-ray elemental mapping of P, F, Cl Geologia Croatica 67/2Geologia Croatica 90 Figure 2: Stratigraphic column of the Al Mahr-1 exploration well showing stratigraphic distribution of planktonic foraminifera (Formation names after SPC – Syrian Petroleum Company). Biozones I (Late Turonian–Early Santonian) II (Middle–Late Santonian) III (Early Campanian) IV Globotruncana plummerae Zone (Early–Middle Campanian) V (Late Campanian) VI Globotruncanella havanensis Zone (Late Campanian) VII Pseudoguembellina palpebra Zone (Late Campanian) VIII Racemiguembelina fructicosa Zone (Late Campanian–Early Maastrichtian) IX Abathomphalus mayaroensis Zone (Late Maastrichtian) Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 91 Figure 3: Stratigraphic column of the Palmyra-1 exploration well showing stratigraphic distribution of planktonic foraminifera (Formation names after SPC – Syrian Petroleum Company). Biozones I (Late Turonian–Early Santonian) II (Middle–Late Santonian) III (Early Campanian) IV Globotruncana plummerae Zone (Early–Middle Campanian) V (Late Campanian) VI Globotruncanella havanensis Zone (Late Campanian) VII Pseudoguembellina palpebra Zone (Late Campanian) VIII Racemiguembelina fructicosa Zone (Late Campanian–Early Maastrichtian) IX Abathomphalus mayaroensis Zone (Late Maastrichtian) Geologia Croatica 67/2Geologia Croatica 92 Figure 4: Correlation between the two exploration wells (Al Mahr-1, Palmyra-1) showing planktonic biozones and lithology. Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 93 and Ca was performed using a FEI Quanta 450 SEM with EDAX TEAM EDS at Smith College (Northampton, Mas- sachusetts, USA). 4. BIOSTRATIGRAPHY Biostratigraphic subdivision of the investigated Upper Cre- taceous successions is based on planktonic foraminifera. Stratigraphic ranges of the identified microfossil assemblages indicate a late Turonian to Maastrichtian age. Stratigraphic relationships between the identified planktonic foraminiferal species are shown in Figs. 2 and 3, whereas the lithostrati- graphic and biostratigraphic biozonation correlation between Al Mahr­1 and Palmyra­1 are presented in Fig. 4. The upper Turonian to lowermost Santonian deposits contain planktonic and benthic foraminiferal assemblages characteristic of this stratigraphic range. These poorly diver- sified microfossil assemblages have equal proportions of small benthic and planktonic foraminifera. The Santonian to lower Campanian strata, on the other hand, are characterized by a moderately diversified microfossil assemblage with in- creased abundance and diversity of planktonic foraminifera, and the absence of nominal taxon/zonal markers. High di- versity microfossil assemblages with a dominance of plank- tonic foraminifera and well-preserved index taxa are present in the upper Campanian to upper Maastrichtian deposits. Previous studies of late Cretaceous planktonic foraminif- era from the Palmyrides only generally indicated the age of the deposits (STANKOVIĆ et al., 2003, 2005; DACER et al., 2007). Biostratigraphy of the upper Turonian to Maas- trichtian successions from the Al Mahr-1 and Palmyra-1 wells is correlated with the existing and well-established plank- tonic foraminiferal zonation for the Tethyan realm (ROBA- SZYNSKI & CARON, 1995; ROBASZYNSKI et al., 2000; PREMOLI SILVA & VERGA, 2004), Fig. 5. Since the zonal markers are very rare or absent, alternative planktonic fo- raminiferal species as well as the whole planktonic fora mi- niferal association enable application of standard biozona- tion (ROBASZYNSKI et al., 1984; CARON, 1985; SLITER, 1989; PREMOLI SILVA & SLITER, 1994; ROBASZYN- SKI & CARON, 1995; ROBASZYNSKI et al., 2000; PRE­ MOLI SILVA & VERGA, 2004; CHACON et al., 2004; HU- BER et al., 2008; SARI, 2009; PETRIZZO et al., 2011; PE REZ­RODRÍGUEZ et al., 2012). Nine biozones have been identified in the upper Turo- nian to Maastrichtian succession: Biozone I, Biozone II, Bi- ozone III, IV Contusotruncana plummerae Zone, Biozone V, VI Globotruncanella havanensis Zone, VII Pseudoguem- belina palpebra Zone, VIII Raceemiguembelina fructicosa Zone, and IX Abathomphalus mayorensis Zone. A list of taxa together with author names and year of publication are pro- vided in the Appendix. All of the diagnostic species and some additional taxa typical of the studied foraminiferal assem- blages are illustrated in Figs. 6–9. Biozone I (Figs. 6A–D) Age. Late Turonian–Early Santonian Interval. Al Mahr­1 (185–210 m, Figs. 2 and 4), Pal­ myra­1 (180–200 m, Figs. 3, 4) Assemblage characteristics. As index taxa were not ob- served, the lowest occurrence (LO) of Contusotruncana for- nicata and Globigerinelloides bollii has been used to define the lower boundary of this Zone. This biozone may corre- spond to the Dicarinella concavata Zone (PREMOLI SILVA & SLITER, 1994; ROBASZYNSKI & CARON, 1995; RO- BASZYNSKI et al., 2000; PREMOLI SILVA & VERGA, 2004; SARI, 2006, 2009). The foraminiferal assemblage of this interval is com- posed of rare non-keeled planktonic foraminifera with a wide stratigraphic range: Archaeoglobigerina blowi, A. cretacea (Figs. 6C, D), Whiteinella balthica, Whiteinella sp., Dicari- nella sp. (Fig. 6A) and Marginotruncana sp. (Fig. 6B). In the middle of the biozone Muricohedbergella holmdelensis and Pseudotextularia nuttalli have their lowest occurrence. The most abundant species in the assemblage are Heterohe- lix reussi, H. moremani, and H. globulosa, comprising 38% of the total planktonic association. Biozone I is also charac- terized by very common Pseudotextularia nuttalli, Murico- hedbergella holmdelensis, Muricohedbergella flandrini, marginotruncanids and whiteinellids. In addition, the follow- ing small calcareous benthic foraminifera are present and account for up to 50% of the total foraminiferal association: Bulimina ovulum, Gyroidinoides globosus, Bulimina sp., Gavellinela sp. (Figs. 9A, B), Lenticulina sp., and Nodosa- ria sp. Lithology and palaeoenvironment. Brownish grey to grey limestone (mudstone/wackestone to foraminiferal wacke stone), dolomitic limestone and marl with equal pro- portions of planktonic and calcareous benthic foraminifera indicate accumulation within outer shelf environments. Biozone II (Figs. 6E–I) Age. Middle–Late Santonian Interval. Al Mahr­1 (150–185 m, Figs. 2 and 4), Pal­ myra­1 (140–180 m, Figs. 3, 4) Assemblage characteristics. Because of the absence of a zonal marker, the lower boundary of this Zone is defined approximately by the LO of Globotruncana linneiana, whe- reas the upper boundary coincides with the disappearance of all dicarinellids and whiteinellids. This biozone may corre- spond to the Dicarinella asymetrica Zone (PREMOLI SILVA & SLITER, 1994; ROBASZYNSKI & CARON, 1995; RO- BASZYNSKI et al., 2000; PREMOLI SILVA & VERGA, 2004; SARI, 2006, 29). The first half of Biozone II is characterized by the LO of several new taxa such as Globotruncana bulloides, G. hilli, G. arca, Hendersonites carinatus, Heterohelix striata, H. punctulata, H. planata, Globigerinelloides prairiehillen- sis and G. subcarinatus. Dominant species in this Zone are Heterohelix globulosa and H. reussi (Fig. 6I), while Pseu- dotextularia nuttalli and Heterohelix punctulata are very common. The following planktonic foraminifera continue from the underlying zone: Archaeoglobigerina blowi (Figs. 6E, F), A. cretacea, Contusotruncana fornicata (Figs. 6G, H), Globigerinelloides bollii, Muricohedbergella holmdel- ensis and Marginotruncana sp. Globotruncana arca, G. bul- loides, and G. linneiana range throughout Biozone II but are Geologia Croatica 67/2Geologia Croatica 94 Figure 5: Proposed biozonation for the Upper Cretaceous succession and stratigraphic range of selected species of the Palmyride region compared with the zonal schemes for Tethys (ROBASZYNSKI and CARON, 1995; PREMOLI SILVA & VERGA, 2004). Timescale was adapted from OGG et al. (2008), and sea- level curve is from HAQ et al. (1987). Grey area indycates the presence of phosphate grains. phate grains and foraminiferal assemblage of Biozone II in- dicate open marine, most probably outer shelf depositional environments. Biozone III (Figs. 6J–N) Age. Early Campanian Interval. Al Mahr­1 (120–150 m, Figs. 2 and 4), Palmyra­1 (115–140 m, Figs. 3, 4) Assemblage characteristics. The main characteristic of the microfossil assemblage is the disappearance of margi- notruncanids at the base of this biozone while Pseudoguem- belina costulata has its lowest occurrence. Rugoglobigerina rugosa first occurs in the middle part of Biozone III, whereas Heterohelix reussi has its highest occurrence (HO). This bi- not present in every sample. In the upper part of this biozone Globotruncanita stuartiformis appears for the first time, while Heterohelix moremani and Muricohedbergella flan- drini have their highest occurrence. The genus Heterohelix is very abundant and diverse (6 species), and comprises a very high percentage (46%) of the planktonic association. Among calcareous benthic foraminifera, common taxa with a wide stratigraphic range include: Bulimina ovulum (Fig. 9D), Praebulimina reussi, P. kickapoensis, Gyroidinoides globosus, Bulimina sp., and Lenticulina sp. Small benthic foraminifera make up to 40% of the microfossil assem- blage. Lithology and palaeoenvironment. Foraminiferal mud- stone/wackestone, marl, dolomitic limestones with phos- Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 95 Figure 6: SEM photomicrographs of selected planktonic foraminifera observed in Biozone I (A–D), Biozone II (E–I), Biozone III, (J–N) and Contusotruncana plummerae Zone (Biozone IV, O, P). A Dicarinella sp., Palmyra-1, interval 190–200 m. B Marginotruncana sp., Al Mahr-1, interval 190–200 m. C, D Archae- oglobigerina cretace (C), and detail of the wall texture (D), Al Mahr-1, interval 185–190 m. E, F Archaeoglobigerina blowi (E), and detail of the wall texture (F), Palmyra-1, interval 160–170 m. G, H Contusotruncana fornicata, Al Mahr-1, interval 155–160 m. I Heterohelix reussi, Palmyra-1, interval 140–150 m. J Het- erohelix globulosa, Palmyra-1, interval 130–140 m. K, L Muricohedbergella holmdelensis, Palmyra-1, interval 120–130 m. M, N Heterohelix striata (M), and detail of the wall texture (N), Al Mahr-1, interval 120–130 m. O, P Contusotruncana plummerae, Palmyra-1, interval 100–110 m. Geologia Croatica 67/2Geologia Croatica 96 ozone may correspond to the Globotruncanita elevata Zone (CARON, 1985; SLITER, 1989; PREMOLI SILVA & SLI­ TER, 1994; ROBASZYNSKI & CARON, 1995; ROBA­ SZYN SKI et al., 2000; PREMOLI SILVA & VERGA, 2004; CHACON et al., 2004). Dominant species in the planktonic foraminiferal assem- blage are Hendersonites carinatus, Heterohelix striata, H. punctulata and Pseudotextularia nuttalli, while Globigeri- nelloides bollii is very common. In addition, the microfossil assemblage contains the following planktonic foraminifera that continue from Biozone II: Archaeoglobigerina cre taca, A. blowi Contusotruncana fornicata, Globotruncana linnei ana, G. bulloides, G. hilli, G. arca, Globotruncanita stu ar ti formis, Heterohelix globulosa (Fig. 6J), H. planata, Globi gerinelloides prairiehillensis, G. subcarinatus, Muricohedbergella holmde- lensis, and Muricohedbergella sp. Spe cimens of the genus Het- erohelix remain diverse and constitute the most abundant group in the microfossil association with 47% of the planktonic fo- raminiferal assemblage. Most of the calcareous benthic fo- raminifera that persisted from the underlying Biozone II, and comprise up to 35% of the microfossil assemblage, include Bulimina ovulum, Buli mina sp. (Fig. 9E), Gyroidinoides glo- bosus, Lenticulina rotulata, Lenticulina sp., and Oridosalis sp. Lithology and palaeoenvironment. Argillaceous lime- stone (mudstone/wackestone) and marl together with micro- fossils of Biozone III suggest an open marine, most probably outer shelf depositional environment. Biozone IV: Contusotruncana plummerae Zone (Figs. 6O, P, 7A, B) Definition. Stratigraphic interval from the LO of Contu- sotruncana plummerae to the LO of Radotruncana calcarata (PETRIZZO et al., 2011) Age. Early to Middle Campanian Interval. Al Mahr­1 (85–120 m, Figs. 2 and 4), Palmyra­1 (80–115 m, Figs. 3, 4) Assemblage characteristics. Beside the LO of the zonal marker Contusotruncana plummerae (Figs. 6O, P) the lower boundary of this biozone is also characterized by the LO of Hendersonites carinatus and by the first occurrence of Lae- viheterohelix glabrans, Muricohedbergella mounmouthensis and Heterohelix navarroensis. The planktonic foraminiferal assemblage is similar to that in Biozone III. However, planktonic foraminifera are more abundant and the overall number of species increased. The dominant species include Heterohelix striata, H. planata, Pseudoguembelina costulata and Pseudotextularia nuttalli. In addition, the assemblage is characterized by common Muri- cohedbergella holmdelensis, M. monmouthensis, Globige- rinelloides bollii (Fig. 7A), G. subcarinatus and Heterohelix globulosa. Other species in this Zone include: Archa eoglo- bigerina blowi (Fig. 7B), A. cretacea, Contu so truncana for- nicata, Globotruncana arca, G. bulloides, G. linneiana, G. hilli, Globotruncanita stuartiformis, Heterohelix punctulata, Globigerinelloides prairiehillensis, Muricohedbergella sp. and Rugoglobigerina rugosa. Representatives of the genus Heterohelix decrease in abundance to 32%, whereas pseudog- uembelinids increase in diversity as well as in abundance and can account for up to 10.5% of the total assemblage. The amount of calcareous benthic foraminifera de- creases to 30% of the assemblage. The most common taxa include Bulimina ovulum, Gavelinella monterelensis (Fig. 9C), Gyroidinoides globosus, Lenticulina rotulata, Lenti- culina sp. (Fig. 9F), Gavelinella sp., Gyroidinoides sp., and Cibicidoides sp. Lithology and palaeoenvironment. Argillaceous lime- stone (wackestone), marl, as well as calcareous marl with abundant planktonic foraminifera (70%) indicate outer shelf to upper bathyal depositional environments. Biozone V (Figs. 7C–G) Age. Late Campanian Interval. Al Mahr­1 (80–85 m, Figs. 2 and 4), Palmyra­1 (75–80 m, Figs. 3, 4) Assemblage characteristics. Since the zonal marker is very rare and generally poorly preserved, the base of Bio- zone V is marked by the LO of Laeviheterohelix dentata, Globotruncanella havanensis and Globotruncanita stuarti. This biozone may correspond to the Radotruncana calcarata Zone (ROBASZYNSKI & CARON, 1995; PREMOLI SIL VA & SLITER, 1994; ROBASZYNSKI et al., 2000; PRE MOLI SILVA & VERGA, 2004; SARI, 2006, 2009; HU- BER et al., 2008). The species Heterohelix labellosa appears in the middle part of Biozone V. The planktonic foraminiferal assemblage of this Biozone is abundant and highly diversified. Most plank- tonic foraminifera persist from the underlying Biozone IV, in- cluding Archaeoglobigerina blowi, A. cretacea, Contuso trun- ca na fornicata, Globotruncana arca (Figs. 7E, F), G. bulloides, G. linneinana, G. hilli, Heterohelix globulosa, H. navarroensis, H. planata, H. punctulata, H. striata, Globigerinelloides bol- lii, G. prairiehillensis (Fig. 7G), G. subcarina tus, Laevihetero- helix glabrans, Muricohedbergella holm delensis, M. moun- mouthensis, Muricohedbergella sp., Pseudo guem belina costu lata, Pseudotextularia nuttalli (Fig. 7C), and Rugoglo- bigerina rugosa (Fig. 7D). The genus Heterohelix constitutes 38% of the assemblage and remains the most abun dant group in the planktonic association. In comparison with previous bi- ozones, the diversity of Biozone V increases and the total num- ber of planktonic foraminifera reaches 27 species. Small benthic foraminifera comprise less than 20% of the assemblage and include Bolivina incrassatea, Gavelinella monterelensis, Gavelinella sp., Gyroidinoides globosus, Len- ticulina rotulata, Lenticulina sp., Neoflabelina reticulata, Serovaina complanata, and Cibicidoides sp. Lithology and palaeoenvironment. Argillaceous lime- stone and calcareous marl with abundant planktonic fo- raminifera (80%) indicate outer shelf to upper bathyal depo- sitional environments. Biozone VI: Globotruncanella havanensis Zone (Figs. 7H–J) Definition. Partial range Zone from the LO of Globotrun- ca nella havanensis to the LO of Pseudoguembelina palpe- bra (HUBER et al., 2008) Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 97 Age. Late Campanian Interval. Al Mahr­1 (70–80 m, Figs. 2 and 4), Palmyra­1 (65–75 m, Figs. 3,4) Assemblage characteristics. The zonal marker Globo- truncanella havanensis (Fig. 7H) is relatively rare, while species Pseudoguembelina excolata, Planoglobulina car- seyae and Rugoglobigerina hexacamerata have their lowest occurrence at the base of the Biozone IV. The following planktonic foraminifera continue from the underlying Biozone V: Archaeoglobigerina blowi, A. cre- tacea, Contusotruncana fornicata, Globotruncana arca, G. buloides, G. linneinana, G. hilli, Globotruncanita stuarti- formis, G. stuarti, Heterohelix globulosa, H. navarroensis (Figs. 7I, J), H. planata, H. punctulata, H. striata, Globige- rinelloides bollii, G. prairiehillensis, G. subcarinatus, Lae- viheterohelix glabrans, L. dentata, Muricohedbergella hol- mdelensis, M. mounmouthensis, Muricohedbergella sp., Pse udo guembelina costulata, Pseudotextularia nuttalli, and Rugoglobigerina rugosa. In the middle of the biozone Glo- botruncanella petaloidea, Rugoglobigerina macrocephala and Guembelitria turrita have their LO. Species of genus Heterohelix remain the most abundant group in the micro- fossil assemblage with 37.5% representation. Globotrunca- nids increase in diversity (8 species) and abundance (13.5%), and become an important component of the planktonic as- semblage. In addition, small benthic foraminifera comprise 15% of the total fauna and include Bolivina incrassata, Ga- veli nella monterelensis, Gavelinella sp., Gyroidinoides glo- bosus, Lenticulina rotulata, Lenticulina sp., Neoflabelina re- ticulata, Serovaina complanata, Cibicidoides sp. (Fig. 9H), and an agglutinated form Spiroplectamina sp. (Fig. 9G). Lithology and palaeoenvironment. Marl and argillaceous limestone (mudstone/wackestone) with abundant planktonic foraminifera (85%) suggest outer shelf to upper bathyal dep- ositional environments. Biozone VII: Pseudoguembelina palpebra Zone (Figs. 7K–P, 8A) Definition. Partial range Zone from the LO of Pseudog- uembelina palpebra to the LO of Racemiguembelina fructi- cosa (HUBER et al., 2008) Age. Late Campanian–Early Maastrichtian Interval. Al Mahr­1 (35–70 m, Figs. 2 and 4), Palmyra­1 (30–65 m, Figs. 3, 4) Assemblage characteristics. Pseudoguembelina palpe- bra (Fig. 7N) is consistently present in this biozone. The first occurrence of Globotruncanella pschadae, Pseudoguembe- lina kempensis and Racemiguembelina powelli is recorded in the lower part of this biozone. Gansserina gansseri (Figs. 7O, P) is present, but very rare, throughout this interval.Very common species in the assemblage include Heterohelix glob- ulosa, Pseudoguembelina costulata (Fig. 7M), Heterohelix striata, and H. navarroensis. Common species include Het- erohelix labellosa, Pseudoguembelina excolata, Pseudotex- tularia nuttalli, and Rugoglobigerina rugosa (Figs. 7L, 8A). Other representative species are Archaeoglobigerina blowi, A. cretacea, Contusotruncana fornicata, Globotruncana arca,G. bulloides, G. linneiana, G. hili, Globotruncanella havanensis, G. petaloidea, Globotruncanita stuarti, G. stu- artiformis, Guembelitria turrita, G. cretacea, Heterohelix planata, H. punctulata, Globigerinelloides bollii, G. prai- riehillensis, G. subcarinatus, Laeviheterohelix dentata, L. glabrans, Muricohedbergella holmdelensis, M. mounmoth- ensis, Muricohedbergella sp., Planoglobulina carseyae, Ru- goglobigerina hexacamerata (Fig. 7K), and R. macroce- pha la. The middle part of Biozone VII is characterized by the lowest occurrence of Abathomphalus intermedius, Glo- botruncanella minuta and Globotruncanita pettersi, while Globigerinelloides bollii become extinct. The upper part of this Biozone is also characterized by the lowest occurrences of Pseudotextularia intermedia and Globotruncanita conica. In comparison with Biozone VI, biodiversity significantly increases throughout Biozone VII and reaches the maximum of 41 species. This increase is partly related to speciation of globotruncanids, (represented by 12 species). The genus Het- erohelix remains the dominant group with 34% abundance, whereas pseudoguembelinids remarkably increase up to 17%. Globotruncanids, despite numerous species, represent 15% of the planktonic foraminiferal population. Among calcareous benthic foraminifera the most com- mon taxa are: Bolivina incrassata, Bolivinoides miliaris, Gavelinella monterelensis, Gavelinella sp., Gyroidinoides globosus, Lenticulina rotulata (Fig. 9J), Lenticulina sp., Ne- oflabelina reticulata (Fig. 9I), Oridorsalis umbonatus, Sero- vaina complanata, Cibicidoides sp., Nodosaria sp., and ag- glutinated Spiroplectammina sp. The proportion of small benthic foraminifera significantly decreases and they make up only 10% of the microfossil assemblage. Lithology and palaeoenvironment. Marl and argillaceous limestone (mudstone/wackestone) with high proportions of planktonic species (up to 90%) imply continuous deepening of this marine realm and deposition in outer shelf to upper bathyal environments. Biozone VIII: Racemiguembelina fructicosa Zone (Figs. 8B–E) Definition. Partial range Zone from the LO of Race mi- guembelina fructicosa to the LO of Abathomphalus ma- yaroensis (HUBER et al., 2008; PEREZ­RODRÍGUEZ et al., 2012) Age. Early Maastrichtian Interval. Al Mahr­1 (30–35 m, Figs. 2 and 4), Palmyra­1 (25–30 m, Figs. 3, 4) Assemblage characteristics. Beside the nominate spe- cies Racemiguembelina fructicosa (Fig. 8E), Planoglobulina acervulinoides and Pseudotextularia elegans are also re- corded for the first time in the lower part of this biozone. In the same horizons Contusotruncana fornicata and C. plum- merae have their HO. Dominant species in the assemblage are Heterohelix globulosa, H. striata and H. navarroensis. Common species include Rugoglobigerina rugosa and R. macrocephala. The planktonic foraminiferal assemblage is similar to that in Biozone VII (41 species) and consists of the following species: Abathomphalus intermedius, Archae- Geologia Croatica 67/2Geologia Croatica 98 Figure 7: SEM photomicrographs of selected planktonic foraminifera observed in Contusotruncana plummerae Zone (Biozone IV, A, B), Biozone V (C–G), Globotruncanella havanensis Zone (Biozone VI, H–J) and Pseudoguembelina palpebra (Biozone VII, K–P). A Globigerinelloides bollii, Al Mahr-1, interval 95– 100 m. B Archaeoglobigerina blowi, Al Mahr-1, interval 85–90 m. C Pseudotextularia nuttalli, Al Mahr-1, interval 80–85 m. D Rugoglobigerina rugosa, Al Mahr-1, interval 80–85 m. E, F Globotruncana arca (E), and detail of the wall texture (F), Palmyra-1, interval 75–80 m. G Globigerinelloides prairiehillensis, Palmyra-1, interval 75–80 m. H Globotruncanella havanensis, Palmyra-1, interval 65–70 m. I, J Heterohelix navarroensis (I), and detail of the wall texture (J), Palmyra-1, interval 60–65 m. K Rugoglobigerina hexacamerata, Al Mahr-1, interval 65–70 m. L Rugoglobigerina rugosa, Al Mahr-1, interval 65–70 m. M Pseu- doguembelina costulata, Al Mahr-1, interval 60–65 m. N Pseudoguembelina palpebra, Palmyra-1, interval 50–60 m. O, P Gansserina gansseri, Al Mahr-1, in- terval 45–50 m (O), and Palmyra-1, interval 40–50 m (P). Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 99 oglobigerina blowi, A. cretacea, Contusotruncana fornicata, Gansserina gansseri, Globotruncana arca, G. bulloides, G. linneiana, G. hilli,Globotruncanella havanensis, G. minuta, G. petaloidea (Fig. 8B), Globotruncanita conica, G. stuarti, G. stuartiformis (Figs. 8C, D), G. pettersi, Guembelitria tur- rita, G. cretacea, Heterohelix labellosa, H. planata, H. punc- tulata, Globigerinelloides bollii, G. prairiehillensis, G. subcarinatus, Laeviheterohelix dentata, L. glabrans, Mu rico- hedbergella holmdelensis, M. mounmothensis, Muricohed- bergella sp., Planoglobulina carseyae, Pseudoguembelina costulata, P. excolata, P. kempensis,P. palpebra, Pseudotex- tularia nuttalli, P. intermedia, Rugoglobigerina hexacam- erata, and Racemiguembelina powelli. Representatives of the genus Heterohelix increase in abundance to 38%, and rugoglobigerinids are more common here than in the under- lying biozones and make up to 14% of the planktonic fo- raminiferal association. Pseudoguembelinids that reached a peak in the previous biozone drop in abundance to 8%, and globotruncanids also decrease in occurrence towards to the top of Biozone VII. Small benthic foraminifera comprise less than 10% of the assemblage and include Bolivina incrassata, Gavelinella monterelensis, Gavelinella sp., Gyroidinoides globosus, Len- ticulina rotulata, Lenticulina sp., Neoflabelina reticulata, Stensioeina pommerana (Figs. 9K, L), Cibicidoides sp., and agglutinated species Gaudryina laevigata. Lithology and palaeoenvironment. Marl and argillaceous limestone (mudstone/wackestone) contain a high proportion of rich and very well preserved planktonic foraminifera sug- gesting deposition in upper to middle bathyal environments. Biozone IX: Abathomphalus mayaroensis Zone (Figs. 9F–P) Definition. Interval Zone from the LO of the nominal taxon to the extinction of most of the Cretaceous planktonic foraminifera (PREMOLI SILVA & SLITER, 1994; ROBA- SZYN SKI & CARON, 1995; ROBASZYNSKI et al., 2000; PREMOLI SILVA & VERGA, 2004; CHACON et al., 2004) Age. Late Maastrichtian Interval. Al Mahr­1 (10–30 m, Figs. 2 and 4), Palmyra­1 (5–25 m, Figs. 3, 4) Assemblage characteristics. The zonal marker Abathom- phalus mayaroensis (Figs. 8F, M, N) is represented by a few specimens. Very common species in the planktonic fora mi- niferal assemblages include Heterohelix globulosa, H. stri- ata, Rugoglobigerina rugosa, Heterohelix navarroensi, H. planata and Muricohedbergella mounmouthensis, while com- mon species include Heterohelix labellosa, Globotruncana arca, Rugoglobigerina macrocephala and Muricohedberg- ella holmdelensis. This very well preserved and highly di- verse assemblage also contains other species such as Glo- botruncana hilli, Globotruncanella havanensis, G. minuta, G. petaloidea, G. pschade, Globotruncanita conica (Fig. 8P), G. stuarti, G. stuartiformis (Figs. 8K, L), G. pettersi, Guem- belitria cretacea, Heterohelix punctulata, Pseudoguembelina excolata (Figs. 8G, H), P. kempensis, P. palpebra, Pseudo- textularia elegans (Figs. 8I, J), P. intermedia, P. nuttalli, Pl- anoglobulina carseyae, and Rugoglobigerina hexacamerata. A large overturn in planktonic fauna occurred within Bio zone IX due to the extinction and disappearance of many species at the base of the biozone, including: Archaeoglobige- rina blowi, A. cretacea, Globotruncana bulloides and Glo- bigerinelloides prairiehillensi.Furthermore, species such as Aba- thom phalus intermedius (Fig. 8O), Gansserina gansseri, Glo botruncana linneiana, Pseudoguembelina costulata and Racemiguembelina powelli become rare and then disappear in the middle part of Biozone IX. Species of the genus Het- erohelix remain the dominant group in the planktonic assem- blage with the same abundance of 36%, whereas globotrun- canids and rugoglobigerinids have almost the same abun dance as in the underlying Biozone VIII. The very high overall di- versity (40 species) of Biozone IX, although somewhat lower than in Biozone VIII, dramatically decreases at the end of the zone when most planktonic foraminiferal species become ex- tinct. Only a few species such as Muricohedbergella holmdel- lensis, M. monmouthensis and Guembelitria cretacea cross the Cretaceous/Palaeogene boundary. The calcareous benthic species Bolivinoides draco (Fig. 9P) is also characteristic of Abathomphalus mayaroensis Zone (GAWOR­BIEDOWA, 1984). In addition to this spe- cies, many other small benthic foraminifera also occur: Bolivina incrassata, Gyroidinoides globosus (Fig. 9N), Len- ticulina rotulata, L. münsteri, Neoflabelina reticulata, Sten- sioeina pommerana, Cibicidoides sp. (Fig. 9M) as well as agglutinated forms Gaudryina laevigata (Fig. 9O) and Spiroplectammina sp. Lithology and palaeoenvironment. Marl, argillaceous limestone (mudstone/wackestone) and slightly dolomitized limestone (foraminiferal wackestone) contain rich and very well preserved foraminiferal assemblages (plankton/benthos ratio is 94:6), suggesting a permanent open­marine influence and deposition in upper to middle bathyal environments. 5. INTERPRETATION AND DISCUSSION This detailed study of the Late Turonian–Maastrichtian plank- tonic and benthic foraminiferal assemblages provides the ba sis for biostratigraphic and palaeoenvironmental interpre- tations of the successions examined. A total of 56 planktonic foraminiferal species belonging to 20 different genera have been identified. Abundant and moderately to highly diverse and generally well preserved planktonic foraminiferal assem- blages enabled biozonation and identification of the follow- ing biozones: Biozone I, Biozone II, Biozone I, IV Contus- otruncana plummerae Zone, Biozone V, VI Globo truncanella ha va nensis Zone, VII Pseudoguembelina palpebra Zone, VIII Racemiguembelina fructicosa Zone and IX Abathom- phalus mayorensis Zone. Identification of possible strati- graphic gaps in the Upper Cretaceous successions examined here was very difficult because the drill cuttings were sam- pled every 5–10 metres. According to BREW (2001), the Upper Cretaceous strata succession of the Palmyride area is characterized by progressively deeper water environments. Evidence for some minor compression and uplift has been documented for the latest Cretaceous of this area, together with an associated minor sedimentary hiatus at the Creta- Geologia Croatica 67/2Geologia Croatica 100 Figure 8: SEM photomicrographs of selected planktonic foraminifera observed in the Pseudoguembelina palpebra Zone (Biozone VII, A) Racemiguembe- lina fructicosa Zone (Biozone VIII, B–E) and Abathomphalus mayaroensis Zone (Biozone IX, F–P). A Rugoglobigerina rugosa, Palmyra-1, interval 40–50 m. B Globotruncanella petaloidea, Al Mahr-1, interval 30–35 m. C, D Globotruncanita stuartiformis (C), and detail of the wall texture (D), Al Mahr-1, interval 30–35 m. E Racemiguembelina fructicosa, Palmyra-1, interval 25–30 m. F Abathomphalus mayaroensis, Al Mahr-1, interval 30–35 m. G, H Pseudoguembelina exco- lata (G), and detail showing wall texture (H), Palmyra-1, interval 20–25 m. I, J Pseudotextularia elegans (I), and detail showing wall texture (J), Palmyra-1, interval 20–25 m. K, L Globotruncanita stuartiformis, Al Mahar-1, interval 20–25 m. M, N Abathomphalus mayaroensis, Al Mahr-1, interval 15–20 m (M), and Palmyra-1, interval 10–20 m (N). O Abathomphalus intermedius, Palmyra-1, interval 10–20 m. P Globotruncanita conica, Palmyra-1, interval 10–20 m. Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 101 Figure 9: SEM photomicrographs of selected benthic foraminifera observed in the upper Turonian–Maastrichtian sequence of the Palmyride Region. A, B Gavelinella sp., Al Mahr-1, interval 185–190 m (A), and Palmyra-1, interval 180–190 m. C Gavelinella monterelensis, Al Mahr-1, interval 115–120 m. D Bulimina ovulum, Al Mahr-1, interval 150–155 m. E Bulimina sp., Palmyra-1, interval 130–140 m. F Lenticulina sp., Palmyra-1, interval 90–100 m. G Spiroplectammina sp., Al Mahr-1, interval 70–80 m. H Cibicidoides sp., Palmyra-1, interval 65–75 m. I Neoflabelina reticulata, Palmyra-1, interval 50–60 m. J Lenticulina rotulata, Al Mahr-1, interval 45–50 m. K, L Stensioeina pommerana, Al Mahr-1, interval 25–30 m. M Cibicidoides sp., Palmyra-1, interval 15–25 m. N Gyroidinoides globosus, Palmyra-1, interval 15–25 m. O Gaudryna laevigata, Al Mahr-1, interval 15–20 m. P Bolivinoides draco, Palmyra-1, interval 10–20 m. Geologia Croatica 67/2Geologia Croatica 102 ceous/Palaeogene boundary (BREW, 2001). To the northeast of the Palmyride area, however, a widespread unconformity has been documented for the Turonian–Coniacian. Accord- ing to BREW (2001), during the Campanian and early Maas- trichtian in the Palmyride area of Syria, progressively deeper water carbonate facies and pelagic marly limestones of the Shiranish formation were deposited. A significant period of Late Cretaceous deformation in northeastern Syria began in the latest Campanian or earliest Maastrichtian (BREW, 2001). The boundary between the Soukhne (massive lime- stone) formation and the syn-extensional Shiranish forma- tion is unconformable, suggesting a major pre-extensional stratigraphic hiatus in that area. The foraminiferal assemblage of Biozone I (late Turo- nian–early Santonian, upper part of the Judea Formation; Figs. 2–5 and 10) is moderately preserved. This biozone may corespond to the Dicarinella concavata Zone, and is char- acterized by the LO of Gobigerinelloides bollii and Contu- sotruncana fornicata, as well as by abundant Heterohelix reussi, H. globulosa and H. moremani. Opportunistic (r-strat- egists) biserial taxa heterohelicids and globular archeoglo- bigerinids (PREMOLI SILVA & SLITER, 1999; PETRIZZO, 2002, 2003) are important components in this biozone. The dominant species Heterohelix globulosa inhabits subsurface levels in the water column (ABRAMOVICH et al., 2003). Domination of opportunistic biserial heterohelicids and other genera with simple morphology (Muricohedbergella, Pseu- dotextularia, Archeoglobigerina and Whiteinella), which comprise up to 87 % of the total planktonic foraminiferal as- semblage, indicate palaeoceanographic conditions favorable for opportunistic (r-strategist) organisms, such as a high pro- ductivity ocean with generally cooler but fluctuating climate, well developed oxygen minimum zone, common up-wellings and cyclic eutrophications of the surface water (BOERSMA & PREMOLI SILVA, 1989; NEDERBRAGT et al., 1998; PETRIZZO, 2002). On the other hand, the occurrence of K-strategists (Di- carinella and Marginotruncana), although present in a smaller percentage in the planktonic assemblage, indicates warm stable episodes with oligotrophic oceanic conditions and well developed water column stratification, which are favourable for these two groups with more complex test ar- chitecture (PETRIZZO, 2002). Almost equal proportions of small benthic and planktonic foraminifera in limestones (mudstone/wackestone to foraminiferal wackestone) and marl suggest deposition in outer shelf environments (OLS- SON & NYONG, 1984; BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005 The most important characteristic of Biozone II (mid- dle–late Santonian, the Soukhne Formation; Figs. 2–5 and 10) is the high level of speciation of planktonic foraminifera. This Biozone is determined by the first appearance of sev- eral new taxa including Globotruncana linneiana, G. arca, G. bulloides, G. hilli, Hendersonites carinatus, Heterohelix planata, H. punctulata, H. striata, Globigerinelloides prai- riehillensis, and Globotruncanita stuartiformis, which may suggest the Dicarinella asymetrica Zone. The foraminiferal assemblage is moderately diverse and better preserved rela- tive to biozone I. Heterohelicids experienced speciation dur- ing this Biozone; their abundance increased to 46.5%, and they remained a dominant group until the end of the Creta- ceous. As opportunistic planktonic foraminifera heteroheli- cids inhabit more nutrient-rich waters and are indicators of cooler and unstable environments (NEDERBRAGT, 1991; NEDERBRAGT et al., 1998; PETRIZZO, 2002). Their spe- ciation is most likely induced by a somewhat cooler but var- iable climate and anoxic events during the middle Santonian. Beside heterohelicids, other small-sized forms with simple test-morphology, such as muriciohedbergellids, archeoglo- bigerinids and globigerinelloids, are very common in the planktonic assemblage. All of these groups belong to oppor- tunistic taxa that have a great reproductive potential in eu- trophic and somewhat mesotrophic environments with a very well developed oxygen minimum layer (NEDERBRAGT, 1991). Small-sized heterohelicids indicate expansion of the oxygen minimum zone (OMZ) due to increased surface wa- ter productivity and depletion of oxygen in subsurface wa- ters by oxidation of organic carbon (LECKIE, 1987; LECKIE et al., 1998; KELLER & PARDO, 2004; PARDO & KEL- LER, 2008; ASHCKENAZI­POLIVODA et al., 2011). Het- erohelicidae were found to be very common in most of the OMZ suggesting high productivity and/or some tolerance to subsurface oxygen depletion (ASHCKENAZI-POLIVODA et al., 2011). In addition, abundant phosphate grains in the upper part of this zone, support the interpretation that Bio- zone II was characterized by high palaeoproductivity, rela- tively constant and high food supply and moderate increase in bottom water aeration. Very high productivity during this biozone was supported by a fluctuating climate and up­well- ing cycles, which brought nutrient-rich water into the envi- ronments inhabited by heterohelicids and upper-middle bath- yal benthic foraminifera. At the upper boundary of Biozone II all dicarinellids and whiteinellids became extinct. The pro- portion of planktonic species increased and reached up to 60% of the microfossil assemblage present in foraminiferal mudstone/wackestone, marl and dolomitic limestones that represent an open marine, most probably outer shelf envi- ronments (OLSSON & NYONG, 1984; BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005). Biozone III (early Campanian, the Soukhne Formation; Figs. 2–5 and 10) is marked by the disappearance of margi- notruncanids in its base and by the LO of Pseudoguembelina costulata and Rugloglobirerina rugosa. This planktonic as- semblage may correspond to the Globotruncanita elevata Zone. Planktonic and benthic foraminiferal assemblages are rich and moderately to well preserved. The proportion of planktonic species reaches up to 65% and indicates further deepening of this realm (BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005; DARVISHZAD & ABDOLALIPOUR, 2009). The most common species are opportunistic (r­strat- egists) taxa: Hendersonites carinatus, Heterohelix punctu- lata, H. striata and Pseudotextularia nuttalli. Although char- acterized by different deposits, i.e., limestone (foraminiferal mudstone/wackestone) and calcareous marl, relative to Bio- zone II, the deposition of these strata continued within the same open marine, probably outer shelf settings. Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 103 Phosphate grains are very common in dolomitic lime- stones from the upper part of Biozone II in Al Mahr-1 (Figs. 2 and 4), and in the uppermost part of Biozone II and the lowermost part of Biozone III in Palmyra­1 (Figs. 3, 4). Abundant phosphate grains generally indicate some very specific geological and palaeoenvironmental conditions, such as oxygen deficiency, upwelling conditions, and trans- gressive intervals (HAQ et al., 1987; REISS, 1988; AL- MOGI­LABIN et al., 1993; WIDMARK & SPEIJER, 1997; JARVIS et al., 2002; PUFHAL et al., 2003; SOUDRY et al., 2006; ASHCKENAZI­POLIVODA et al., 2011). It is pos- sible that such palaeoceanographic conditions, especially up- welling, increased food supply and primary production in the surface and subsurface marine environments, and thus also indirectly affected higher production and domination of oportunistic (r-strategists) planktonic foraminiferal species during Biozones II and III. The lowest occurrence of Laeviheterohelix glabrans and Muricohedbergella mounmouthensis and rare Contusotrun- cana plummerae in the planktonic foraminiferal assemblage of Biozone IV (middle–late Campanian, the Shiranish For- mation; Figs. 2–5 and 10) suggest the Contusotruncana plummerae Zone. This biozone has been appointed by PETRIZZO et al. (2011) for the lower–middle Campanian of tropical and subtropical areas because of the difficulties in using the first occurrence datum of Globotruncana ven- tricosa in low latitude successions from the Tethyan Realm. Species of the genus Heterohelix dominated in the previous Biozone III but decrease to 32.5% in Biozone IV, whereas the abundance of two genera Muricohedbergella and Pseu- doguembelina significantly increases up to 15% and 10.5%, respectively. Although the opportunistic (r-strategists) spe- cies are still dominant, the specialized taxa (K-strategists) such as globotruncanids (PREMOLI SILVA & SLITER, 1999; PETRIZZO, 2002; DUBICKA & PERYT, 2012) in- crease in the overall number of species and also slightly in- crease in abundance within this planktonic assemblage. This indicates mesotrophic to more oligotrophic environmental conditions that are favorable for keeled globotruncanids. The well-preserved foraminiferal assemblage and high propor- tion of planktonic foraminifera (70%) in the argillaceous limestones marl and calcareous marls of Biozone IV suggest an open marine, probably outer shelf to upper bathyal envi- ronment (BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005). The lowest occurrence of planktonic foraminifera Lae- viheterohelix dentata, Globotruncanella havanensis and Globotruncanita stuarti in Biozone V (late Campanian, the Shiranish Formation; Figs. 2–5 and 10) may suggest the Ra- dotruncana calcarata Zone. The well-preserved foraminif- eral assemblage is rich, diverse and has an increased number of K-strategists (keeled globotruncanids; PREMOLI SILVA & SLITER, 1999). Globotruncanids have a complex test morphology and usually inhabit the subsubsurface water col- umn (PETRIZZO, 2002; ABRAMOVICH et al., 2003; DU- BICKA & PERYT, 2012), which implies good stratification of the water column and a tropical to subtropical climate dur- ing the late Campanian (LECKIE, 1989; HUBER et al., 1995; DARVISHZAD & ABDOLALIPOUR, 2009). Depos- its and sedimentary settings remain the same as for Biozone IV, i.e., deep-water marine environments. The zonal marker Globotruncanella havanensis is rela- tively rare in Biozone VI (late Campanian, the Shiranish Formation; Figs. 2–5 and 10), and the base of this Biozone is indicated by the LO of Pseudoguembelina excolata, Pla- noglobulina carseyae and Rugoglobigerina hexacamerata. The foraminiferal assemblage is rich and well preserved. Al- though the opportunistic group heterohelicids stay dominant group in the planktonic assemblage with 37.5% abundance, K-strategists, such as keeled globotruncanids, become an important component in the planktonic foraminiferal assem- blage with 9 species and 13.5% abundance. These point to stable environmental conditions, such as an oligotrophic ocean with a tropical to subtropical climate, well stratified water column, stable thermocline and other favourable pal- aeoceanographic parameters for r/K and K-selected group of planktonic foraminifera (LECKIE, 1989; MURRAY, 1991; DARVISHZAD & ABDOLALIPOUR, 2009). Abun- dance of species from the genus Globigerinelloides and Mu- ricohedbergella (r-selected forms) show inverse trends in comparison with the previous biozones and decrease to 7.5% and 6%, respectively (Fig. 10). The proportion of planktonic species increased and reached up to 85% of the microfossil assemblage present in marl and argillaceous limestone (mud- stone/wackestone) that represent an open marine, outer shelf to upper bathyal environments (OLSSON & NYONG, 1984; BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005). The Pseudoguembelina palpebra Zone (Biozone VII, late Campanian–early Maastrichtian, the Shiranish Forma- tion; Figs. 2–5 and 10) is characterized by the LO of P. palpe- bra, which is consistently found throughout the interval in moderate abundance. Also, the lowest occurrence of Glo- botruncanella pschadae, Pseudoguembelina kempensis and Racemiguembelina powelli is recorded in the lower part of this biozone. Gansserina gansseri is very rare and poorly preserved in the investigated samples, and therefore P. palpe- bra serves as a better zonal marker for the uppermost Cam- panian, as also reported by HUBER at al. (2008) from sub- tropical North Atlantic (Blake Nose). The planktonic foraminiferal assemblage of Biozone VII is rich, very well preserved and in comparison with Biozone VI, biodiversity throughout this interval significantly increases (to 41 spe- cies). Opportunistic representatives of the genus Heterohelix are still the dominant group in the planktonic assemblage with 34% abundance. Also, the genus Pseudoguembelina, known as a successful surface and subsurface dweller in tropical and subtropical open ocean (NEDERBRAGT, 1989; HUBER, 1992; ABRAMOVICH et al., 2003), significantly increased in abundance up to17%. Species of the genus Pseu- doguembelina are strongly photosymbiotic and their expan- sion is related to favourable palaeoecological conditions in the Late Cretaceous ocean, such as the presence of warm and oligotrophic surface ocean waters (D´HONDT & ZACHOS, 1998; ABRAMOVICH et al., 2003). The proportion of planktonic foraminifera accounts for up to 90% of the mi- crofossil assemblages found in marl and argillaceous lime- Geologia Croatica 67/2Geologia Croatica 104 stone (mudstone/wackestone), which implies further deep- ening of this realm and deposition in outer shelf to upper bathyal environments (BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005; DARVISHZAD & ABDOLALIPOUR, 2009). The lowest occurrence of planktonic foraminifera Pla- noglobulina acervulinoides and Peudotextularia elegans in association with the rare zonal marker Racemiguembelina fructicosa, is indicative of Biozone VIII (early–late Maas- trichtian, the Shiranish Formation; Figs. 2–5 and 10). This biozone is characterised by a diverse and very well preserved planktonic foraminiferal assemblage with 41 species, similar to that from Biozone VII. Rugoglobigerinids increased in abundance and reached up to 14% of the assemblage, while representatives of the genus Heterohelix still dominate the assemblage with 38%. Species of the genus Rugoglobigerina inhabit surface and subsurface habitats (ABRAMOVICH et al., 2003), and are known as symbiotic organisms (D´HONDT & ZACHOS, 1998). Speciation of planktonic foraminifera in this biozone is likely supported by good water column stratification and opening of new niches favourable for all groups of planktonic foraminifera. Many subsurface dwell- ers, such as several species of the genus Globotruncana, ac- quired adaptation to the thermocline habitat (ABRAMOV- ICH et al., 2003). All of these changes correspond very well to the documented global sea level fluctuations and alternat- ing cooler and warmer periods in the early to late Maastrich- tian (HAQ et al., 1987; Fig. 5). During Zone VIII the sedi- mentary setting was a deep sea environment (upper bathyal), as a result of further deepening of this sedimentary realm. The planktonic foraminiferal assemblage of the latest Cretaceous Abathomphalus mayaroensis Zone (Biozone IX, late Maastrichtian, the Shiranish Formation; Figs. 2–5 and 10) is very similar to those in Biozone VIII and is rich in the overall number of species (39) as well as in the number of individuals. An important characteristic of this biozone is the increased number of K-strategist species of globotrun- canids (15.5%), and planoglobulinids (4%), whereas rug- oglobigerinids retained their abundance (14%). Some op- portunistic species of the genus Muricohedbergella also show an increase and constitute 9% of the assemblage, while members of the genus Globigerinelloides (2%) decrease in abundance (Fig. 10). Species of the genus Heterohelix have an almost equal abundance (36%) compared to Biozone VIII. Composition of the planktonic foraminiferal assemblage with a significantly higher percentage of K­strategist special- ists, which inhabit subsurface and thermocline layers, indi- cates an oligotrophic ocean with a very well stratified water column supported by a stable thermocline (PREMOLI SILVA & SLITER, 1999; PEARSON et al., 2001; ABRAM- OVICH et al., 2003). On the other hand, the symbiont­bear- ing taxa Rugoglobigerina, Pseudoguembelina and Hetero- helix were very well adapted to surface and subsurface oligotrophic ocean water (D´HONDT & ZACHOS, 1998; PEARSON et al., 2001; ABRAMOVICH et al., 2003). In the upper part of Biozone IX, the decreased numbers of glo- botruncanid species indicate fluctuating climate, sea­level changes and up-welling cycles which could cause instability in the water column and unfavourable environmental condi- tions for K­strategists (ABRAMOVICH & KELLER, 2002; HAQ et al., 1987). This high diversity abruptly decreases at the end of the Biozone, when most planktonic foraminiferal species became extinct. Marl, argillaceous limestone (mud- stone/wackestone), and slightly dolomitized limestone (fo- raminiferal wackestone) contain rich and very well preserved foraminiferal assemblages, whereas planktonic foraminifera reach over 90% of the entire community suggesting deposi- tion in upper bathyal environments (BOERSMA, 1988; MURRAY, 1991; GRÄFE, 2005; DARVISHZAD & AB- DOLALIPOUR, 2009). All these facts indicate the Late Cre- taceous as being a long, warm and relatively stable period with palaeoceanographic conditions favourable for all groups of planktonic foraminifera which inhabited different niches in a well stratified water column (LECKIE, 1989; HUBER et al., 1995; PREMOLI SILVA & SLITER, 1999; PETRIZZO, 2002, PEARSON et al., 2001; ABRAMOVICH et al., 2003; DUBICKA & PERYT, 2012). The Soukhne Formation (Santonian–Early Campanian, Biozones II and III, Figs. 2–4) contains phosphate grains. Similar phosphate deposits are widespread in many parts of the Levantin region (i.e., Israel, Jordan, Iraq, Turkey and Egypt; AL MALEH & MOUTY, 1994; PUFHAL et al., 2003; ABED at al., 2005; BAIOUMY & TADA, 2005; SOUDRY et al., 2006; ASHCKENAZI­POLIVODA et al., 2011; SCH- NEIDER­MOR et al., 2012). The phosphate deposits in Syria formed in response to a high-productivity upwelling regime that persisted on the southern margins of the Tethys during the Late Cretaceous (AL MALEH & MOUTY, 1994). The planktonic assemblages associated with phosphate grains in the Palmyride strata are characterized by domination of op- portunistic (r-strategists) taxa such as the genus Heterohelix (Biozone II and III, Figs. 2, 3), which indicates a highly pro- ductive photic zone (REISS, 1988; ALMOGI-LABIN et al., 1993; WIDMARK & SPEIJER, 1997; PUFHAL et al., 2003) and a low oxygen environment (ASHCKENAZI-POLI- VO DA et al., 2011). The benthic assemblages found with phosphates have abundant specimens of the genus Bulimina, which are commonly documented from highly productive, low-oxygen settings from around the world, including for example South America, Morocco, Egypt, Jordan, Iraq and Israel (PUFHAL et al., 2003; ASHCKENAZI­POLIVODA et al., 2011). An additional factor that contributed to the for- mation of phosphate was the enrichment in phosphorous from P-rich deep waters that upwelled in the Palmyrida Ba- sin by currents flowing along the northern edge of the Ara- bian platform (SOUDRY et al., 2006). Warming of the up- welled water and the abundance of nutrients caused the proliferation of plankton, which assimilated, stored and con- centrated phosphate. After the deposition of plankton, a large amount of phosphate dissolved and became concentrated in the sea­floor sediments (AL MALEH & MOUTY, 1994). Such high primary productivity and sea floor phospho- genesis prevailed mainly on the southeastern Tethyan mar- gins as a result of persistent upwelling circulation that recy- cled dissolved phosphorous from the intermediate-depth waters and distributed it to the photic zone (SOUDRY et al., Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 105 2006). The phosphates developed during transgressive peri- ods that promoted carbonate sediment starvation. Simple (in- ternally homogenous) P2O5-enriched phosphate nodules probably replaced calcite nodule precursors in suboxic con- ditions as a result of processes that involved oceanic up- welling, exhumation and burial coupled with alternating oxic and suboxic conditions (MARSHALL-NEILL & RUFFELL, 2004). The presence of phosphate is an important indicator of oxygen deficiency, upwelling conditions, transgressive intervals, and omission surfaces (TRAPPE, 2001), and any future studies of this Cretaceous stratigraphic interval should also include detailed analyses of the associated phosphate grains. 6. CONCLUSIONS The stratigraphic analysis of the Upper Cretaceous strata from the Palmyride area in Syria included a detailed micro- palaeontological investigation of foraminferal assemblages and rock types obtained from drill cuttings in two deep ex- ploration wells (Al Mahr-1 and Palmyra-1). Nine biozones have been identified in the upper Turo- nian to Maastrichtian succession: Biozone I, Biozone II, Bi- ozone III, IV Contusotruncana plummerae Zone, Biozone V, VI Globotruncanella havanensis Zone, VII Pseudoguem- belina palpebra Zone, VIII Raceemiguembelina fructicosa Zone, and IX Abathomphalus mayorensis Zone. During the late Turonian to early Campanian (Biozone I to III) in the investigated Palmyride strata, domination of opportunistic taxa (Heterohelix, Globigerinelloides, Archae- oglobigerina, and Muricohedbergella) implies upwelling, low oxygen conditions and subtropical climate. On the other hand, the rich and highly diversified planktonic assemblages (Biozone IV to IX) with many K-selected taxa (i.e., orna- mented keeled globotruncanids, rugoglobigerinids, pla- noglobulinids, pseudoguembelinids) indicate a tropical to subtropical climate and well­stratified water column during the late Campanian and into the Maastrichtian. In the upper part of Biozone IX, the decreased numbers of globotrunca- nid species indicate a less stratified water column and unfa- vourable environmental conditions for K-strategists. A dra- matic faunal turnover at the end of this Biozone is marked by the extinction of most planktonic foraminifera, with only a few species (e.g., Muricohedbergella holmdelensis, M. montmouthensis and Guembelitria cretacea) present across the Cretaceous/Palaeogene boundary. Phosphate grains are very common in dolomitic lime- stone of the upper part of Biozone II in Al Mahr-1 and in the uppermost part of Biozone II and lowermost part of Biozone III in Palmyra-1 (Soukhne Formation). The phosphate oc- currence helps improve the late Santonian–early Campanian stratigraphic interpretation of this interval because similar deposits occur during this time period elsewhere along the southeastern margins of Tethys (Israel, Jordan, Iraq, Turkey and Egypt). The presence of phosphate in the study area in- dicates oceanic upwelling that caused increased food supply and influenced higher primary marine production, and thus indirectly affected higher production and domination of oportunistic planktonic foraminifera in Biozones II and III. ACKNOWLEDGMENT We thank Bosiljka GLUMAC (Smith College) whose com- ments and suggestions improved this paper, Morana HER- NITZ­KUČENJAK (INA­industrija nafte d.d.) for discus- sion and tehnical support, Robert KOŠĆAL (University of Zagreb) for technical assistance, Renata SLAVKOVIĆ (INA-industrija nafte d.d.) for SEM photomicrografs of for- aminifera, Vladimir VESELI and Ivan A. MESIĆ (INA­in- dustrija nafte d.d.) for sharing well report data. We are grate- ful to Brian T. HUBER (Smithsonian Institution) and Bilal SARI (Dokuz Eylül University) for carefully reviewing the Figure 10: Percentage of Late Cretaceous genera in the planktonic foraminiferal assemblages from the study area in Syria. 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Manuscript received December 06, 2013 Revised manuscript accepted March 08, 2014 Available online June17, 2014 Pecimotika et al.: Planktonic foraminiferal biostratigraphy and paleoecology of Upper Cretaceous deposits from the Palmyride Region, Syria Geologia Croatica 109 APPENDIX 1 List of taxa cited in the text and figure explanations, in al- phabetical order according to genus Planktonic foraminifera Abathomphalus intermedius (BOLLI, 1951) Abathomphalus mayaroensis (BOLLI, 1951) Archaeoglobigerina blowi PESSAGNO, 1967 Archaeoglobigerina cretacea (D’ORBIGNY, 1840) Contusotruncana fornicata (PLUMMER, 1931) Contusotruncana plummerae (GANDOLFI, 1955) Dicarinella sp. Gansserina gansseri (BOLLI, 1951) Gglobigerinelloides bollii (PESSAGNO, 1967) Gobigerinelloides prairiehillensis (PESSAGNO, 1967) Globigerinelloides subcarinatus (BRÖENNIMANN, 1952) Globotruncana arca (CUSHMAN, 1926) Globotruncana bulloides VOGLER, 1941 Globotruncana hilli PESSAGNO, 1967 Globotruncana linneiana (D’ORBIGNY, 1839) Globotruncanella havanensis (VOORWIJK, 1937) Globotruncanella minuta CARON & GONZALEZ DO- NOSO, 1984 Globotruncanella petaloidea (GANDOLFI, 1955) Globotruncanella pschadae (KELLER, 1946) Globotruncanita conica (WHITE, 1928) Globotruncanita pettersi (GANDOLFI, 1955) Globotruncanita stuarti (DE LAPPARENT, 1918) Globotruncanita stuartiformis (DALBIEZ, 1955) Guembelitria cretacea CUSHMAN, 1933 Guembelitria turrita NEDERBRAGT, 1990 Hendersonites carinatus (CUSHMAN, 1938) Heterohelix globulosa (EHRENBERG, 1840) Heterohelix labellosa NEDERBRAGT, 1990 Heterohelix navarroensis LOEBLICH, 1951 Heterohelix moremani (CUSHMAN, 1938) Heterohelix planata (CUSHMAN, 1938) Heterohelix punctulata (CUSHMAN, 1938) Heterohelix reussi (CUSHMAN, 1938) Heterohelix striata (EHRENBERG, 1840) Laeviheterohelix dentata (STENESTAD, 1968) Laeviheterohelix glabrans (CUSHMAN, 1938) Marginotruncana sp. Muricohedbergella flandrini (PORTHAULT, 1970) Muricohedbergeela holmdelensis (OLSSON, 1964) Muricohedbergella mounmouthensis (OLSSON, 1960) Muricohedbergella sp. Planoglobulina carseyae (PLUMMER, 1931) Planoglobulina acervulinoides (EGGER, 1899) Pseudoguembelina costulata (CUSHMAN, 1938) Pseudoguembelina excolata (CUSHMAN, 1926) Pseudoguembelina kempensis ESKER, 1968 Pseudoguembelina palpebra BRÖENNIMANN & BROWN, 1953 Pseudotextularia elegans (RZEHAK, 1891) Pseudotextularia intermedia DE KLASZ, 1953 Pseudotextularia nuttalli (VORWIJK, 1937) Racemiguembelina fructicosa (EGGER, 1902) Racemiguembelina powelli SMITH & PESSAGNO, 1973 Rugoglobigerina macrocephala BRÖENNIMANN, 1952 Rugoglobigerina hexacamerata BRÖENNIMANN, 1952 Rugoglobigerina rugosa (PLUMMER, 1926) Whiteinella balthica DOUGLAS & RANKIN, 1969 Whiteinella sp. Benthic foraminifera Bolivinoides draco (MARSSON, 1878) Bolivinoides miliaris HILTERMANN & KOCH, 1950 Bolivina incrassata REUSS, 1851 Bulimina ovulum REUSS, 1844 Bulimina sp. Cibicidoides sp. Gaudryina laevigata FRANKE, 1914 Gavelinella monterelensis (MARIE, 1941) Gavelinella sp. Gyroidinoides globosus HAGENOW, 1842 Gyroidinoides sp. Lenticulina rotulata (LAMARCK, 1804) Lenticulina münsteri (ROEMER, 1839) Lenticulina sp. Neoflabelina reticulata (REUSS, 1851) Nodosaria sp. Oridorsalis umbonatus (REUSS, 1851) Oridorsalis sp. Praebulimina reussi (MORROW, 1934) Praebulimina kickapoensis (COLE, 1938) Serovaina complanata (CUSHMAN & STAINFORTH, 1945) Spiroplectammina sp. Stensioeina pommerana BROTZEN, 1936 Geologia Croatica 67/2Geologia Croatica 110 APPENDIX 2 Distribution of Late Cretaceous genera of planktonic foraminiferal assemblage from Al Mahr-1 and Palmyra-1 wells. DEPTH (m) 180-185 145-150 125-130 90-95 80-85 65-70 40-45 25-30 20-25 BIOZONE I II III IV V VI VII VIII IX SPECIES RICNESS 14 22 22 24 27 32 41 41 40 Abathomphalus intermedius 1 2 1 Abathomphalus mayaroensis 1 Archaeoglobigerina blowi 15 16 10 6 6 3 3 2 Archaeoglobigerina cretacea 12 11 5 2 3 2 2 1 Contusotruncana fornicata 17 14 5 6 7 4 2 Contusotruncana plummerae 2 6 2 3 Dicarinella sp. 9 5 Gansserina gansseri 2 1 2 Gglobigerinelloides bollii 11 17 27 19 15 8 Gobigerinelloides prairiehillensis 11 15 9 13 7 12 9 Globigerinelloides subcarinatus 9 8 14 9 9 8 6 7 Globotruncana arca 4 4 5 4 9 6 8 14 Globotruncana bulloides 3 3 3 4 2 2 1 Globotruncana hilli 3 3 2 3 4 2 4 11 Globotruncana linneiana 2 2 3 3 4 7 6 Globotruncanella havanensis 10 6 5 7 Globotruncanella minuta 1 1 3 Globotruncanella petaloidea 5 3 3 3 Globotruncanella pschadae 3 3 2 Globotruncanita conica 1 2 1 Globotruncanita pettersi 3 6 3 Globotruncanita stuarti 3 4 2 6 2 Globotruncanita stuartiformis 7 8 7 6 6 9 7 4 Guembelitria cretacea 1 1 Hendersonites carinatus 17 34 Heterohelix globulosa 38 33 19 14 32 30 29 34 29 Heterohelix labellosa 11 14 14 12 14 Heterohelix moremani 31 Heterohelix navarroensis 8 17 27 23 31 21 Heterohelix planata 14 14 32 8 7 6 12 17 Heterohelix punctulata 21 33 9 16 9 9 7 11 Heterohelix reussi 35 31 18 Heterohelix striata 19 35 34 29 32 25 32 27 Laeviheterohelix dentata 5 6 10 4 6 Laeviheterohelix glabrans 6 4 8 7 7 5 Marginotruncana sp. 11 4 Muricohedbergella flandrini 18 Muricohedbergeela holmdelensis 25 19 9 21 18 8 13 7 12 M. mounmouthensis 17 15 7 4 11 17 Muricohedbergella sp. 7 6 7 5 4 Planoglobulina carseyae 7 6 5 8 Pseudoguembelina costulata 15 31 26 23 26 7 4 Pseudoguembelina excolata 6 9 8 7 Pseudoguembelina kempensis 3 4 6 Pseudoguembelina palpebra 13 7 7 Pseudotextularia elegans 11 11 Pseudotextularia intermedia 2 1 1 Pseudotextularia nuttalli 30 23 33 30 16 21 14 6 4 Racemiguembelina fructicosa 1 1 Racemiguembelina powelli 2 2 1 Rugoglobigerina hexacamerata 4 5 9 10 Rugoglobigerina macrocephala 5 15 12 Rugoglobigerina rugosa 16 11 17 21 12 24 26 Whiteinella balthica 11 Whiteinella sp. 9 (%) PLANKT. FORAM. 51.22 59.30 64.27 70.21 80.34 85.17 91.24 92.56 93.98 (%) BENT. FORAM. 48.78 40.67 35.72 29.79 19.66 15.05 8.76 7.44 6.02