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, DECEMBER 2014, 65, 6, 451—470 doi: 10.1515/geoca-2015-0005
Oligocene—Early Miocene planktonic microbiostratigraphy
and paleoenvironments of the South Slovakian Basin
(Lučenec Depression)
SILVIA OZDÍNOVÁ
1
and JÁN SOTÁK
2,3
1
Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, 840 05 Bratislava, Slovak Republic; geolsisa@savba.sk
2
Geological Institute, Slovak Academy of Sciences, Branch: Ďumbierska 1, 974 11 Banská Bystrica, Slovak Republic; sotak@savbb.sk
3
Department of Geography, Faculty of Education, KU Ružomberok, Hrabkovská cesta 1, 03 401 Ružomberok, Slovak Republic
(Manuscript received May 16, 2013; accepted in revised form October 7, 2014)
Abstract: Oligocene and Lower Miocene sediments of the Lučenec Depression were studied to demonstrate the plank-
tonic bioevents and climatic proxies from the Číž Formation (Rupelian) and Lučenec Formation (Chattian—Aquitanian)
on the basis of quantitative analyses of foraminifers and calcareous nannofossils. The oldest nannofossil assemblages of
the Číž Formation belonged to the NP23 Zone and were dominated by Reticulofenestra ornata known for preference of
temperate eutrophic water conditions. An increase in bioproductivity was documented by abundant large-sized plank-
tonic foraminifers (e.g. Turborotalia ampliapertura, Paragloborotalia nana, Subbotina gortanii) and epifaunal to shal-
low-infaunal benthic species. The middle part of the Číž Formation reveals a lowstand progradation of deltaic sediments
of the Rapovce Member. There, the planktonic foraminifers are impoverished in both size and diversity, containing
mostly tenuitellid and cassigerinellid species, probably as a result of decreased salinity and increased anoxia in the Tard
Clay. Contrary of this, the benthic foraminifers are rich, mainly the infaunal forms of uvigerinid species. They probably
proliferated due to a high organic flux from riverine input. Open marine conditions were restored in the upper part of the
Číž Formation above the lowest occurrence (LO) of Cyclicargolithus abisectus on the NP23—NP24 zone boundary. The
transitional interval between the Číž and Lučenec formations (O5/O6 – NP24/25) was approximated by the HOs of
Paragloborotalia opima and Sphenolithus distentus and the LOs of Globigerinoides primordius and Pontosphaera
enormis. Benthic foraminifera of the Lučenec Formation indicate a high productivity and oxygen-deplected environ-
ments. The Oligocene—Miocene boundary in the Lučenec Formation was appointed by the HOs of Helicospahera recta
and Dictyococcites bisectus. Foraminiferal markers of this boundary were established from the HO of Globigerina
ciperoensis and the LO of G. ottnangiensis. The highest nannofossil dating in the Lučenec Formation is recorded by the
LOs of Helicosphaera mediterranea (NN1 Zone) and Discoaster druggi (NN2 Zone). The uppermost part of the Lučenec
Formation contained many Paratethys benthic foraminifera, such as Uvigerina posthantkeni.
Key words: Oligocene—Lower Miocene, calcareous nannofossils, planktonic foraminifers, biozonation, climatic proxies,
paleoecology.
Introduction
The Paleogene formations of the Intra-Carpathian area belong
to two different basins – the Hungarian Paleogene Basin and
Central-Carpathian Paleogene Basin. These basins overlay the
Mesozoic basement nappe units, which were emerged during
the post-Gosau time and differ in type of sedimentation. Shal-
low-water epicontinental deposits fill the Hungarian Paleogene
Basin, while deep-water flysch is characteristic of the Central-
Carpathian Paleogene Basin (Nagymarosy 1990). The two ba-
sins also differ in their geotectonic position. The Hungarian
Paleogene Basin was probably formed as a retro-arc foredeep
basin (Tari et al. 1993), but the Central-Carpathian Paleogene
Basin is interpreted as a fore-arc basin (Soták et al. 2001).
The Hungarian Paleogene Basin extends northward into
the South Slovakian Basin (Fig. 1). Its north-western margin
occurs on the Šahy Antiform, which represents a strike-slip
elevation bordering the Buda Line (Vass 2003). Sediments
of the Hungarian Paleogene Basin occur in the Štúrovo area
and in the Ipe , Lučenec and Rimava depressions. The Paleo-
gene formations were drilled by a series of boreholes such as
Muž a-4 (Seneš 1960; Brestenská & Lehotayová 1960), Cha-
nava LR-9 (Šutovská 1987), Lučenec LC-2 (Vass et al.
2004), Rapovce LR-3 (Vass & Elečko 1992), Číž C-2 (Vass
& Elečko 1982), Muž a LKŠ-1 (Zlinská & Šutovská 1990),
Ivanice FGRK-1 (Zlinská 2009), Bátka RKZ-1 (Vass
1995a), Gemerská Párnica (Halásová et al. 1996), Gemerček
RAO-5 (Nagy et al. 2004) and others (see Vass et al. 2007).
The core of the Rapovce GTL-2 borehole drilled for the
geothermal survey by Geospectrum Ltd. Bratislava in 2007
provided a new opportunity to study the Oligocene and Lower
Miocene formations of the Lučenec Depression. The GTL-2
core provided the reference section for study and under-
standing of the high-resolution stratigraphy and paleoenvi-
ronmental proxies of the Oligocene and Lower Miocene
deposits in the Lučenec Depression.
Geological setting
The Hungarian Paleogene Basin began to develop after a
long-term uplift, lateritic weathering and karst bauxite depo-
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sition in Bakony and the Transdanubian area (Báldi 1986).
The Lutetian transgression gradually flooded lagoons, where
coal-bearing layers were accumulated (Dorog Coal, Obid
Member). Progradation of a shallow-water carbonate plat-
form is documented by the Szöc and Szépvölgy limestones.
Their outer shelf equivalent is represented by the Padrag and
Buda Marls (Báldi-Beke & Báldi 1985; Kázmér et al. 2003).
Late Eocene depocenters were shifted to the Buda Basin,
which rapidly subsided during the Oligocene, until bathyal
conditions were reached. Basin differentiation led to initial
isolation recorded by euxinic facies of the Tard Clay (Báldi
1986). The Rupelian-Chattian sequences of the North Buda
Basin are formed by the Kiscell Clay, Széczény Schlier and
Eger Sandstone (Báldi 1986). The Oligocene transgression is
also expressed by the initial flooding of the basinal area in
Southern Slovakia (Vass 1995a).
The Oligocene sequence of the Lučenec Depression (Fig. 2)
started by the continental fluvial deposits of the Skálnik Mem-
ber, followed by transgressive sediments of the Číž Formation
represented by conglomerates, breccias, sandstones and silt-
stones with marine fauna. The lower members of the Číž For-
mation consist of lagoonal sediments of the Blh Member,
nearshore sediments of the Hostišovce Member and offshore
and carbonate peri-platform facies named the Bátka Lime-
stone. The major part of the Číž Formation belongs to the
Lenártovce Member, which is characterized by deep-water fa-
cies of grey claystones, siltstones and sandstones known as
Kiscell Clay. The regressive sequence of the Číž Formation is
developed in deltaic sediments of the Rapovce Member, rep-
resented by grey sandstones and siltstones with plant rem-
nants, coal laminae and shallow water fauna (Vass 1995a).
The Egerian cycle of deposition in the South Slovakian
Basin is represented by the Lučenec Formation, reaching
thicknesses of more than 1000 m in some places. The sedi-
ments of the Lučenec Formation superposed the deep-water
sediments of the Číž Formation or covered the pre-Tertiary
basement. These transgressive deposits are formed by brec-
cias, conglomerates and sandstones of the Panica Member,
bioclastic limestones and calcareous sandstones with Mio-
gypsina and Lepidocyclina of the Budikovany Member and
clastic limestones, breccias and conglomerates with large
pectinids of the Bretka Member. The Lučenec Formation
is mostly composed of grey calcareous siltstones of the
Szecsény Schlier, which documents the outer shelf up to
upper bathyal environment. Upper Eggerian sediments of
the Lučenec Formation belong to the Opatová Member
(Šutovská-Holcová et al. 1993), representing delta-type sedi-
ments, such as channel gravels, prodelta claystones, delta-
front sandstones and coal seams, developed in a great
thickness (150 m). These deltaic deposits represent the ter-
minal sediments of the North Buda Basin, which evolved
into the Fi akovo/Pétervására Basin from the beginning of
the Early Miocene and then from the Ottnangian into the
Nógrád/Novohrad Basin.
Transgressive sediments of the Fi akovo/Pétervására Basin
are represented by the Fi akovo Formation. They are shallow-
Fig. 1.
Regional geological context of area studied (A) within the Hungarian Paleogene Basin (B – scheme after Nagymarosy 1990), South
Slovakian North Hungarian units (C – sketch map after Vass 1995b) and location of the Rapovce GTL-2 borehole. Legend to sketch
map: 1 – pre-Teriary units; 2 – Hungarian Paleogene Basin; 3, 4 – Fi akovo-Pétervására Basin: 3 – Fi akovo Formation and Péter-
vására Sandstone, 4 – Upper Szécsény and Putnok Schliers; 5 – Eggenburgian rhyodacide tuffs in Gyulakeszi and Bukovina Formations;
6 – Nógrad-Novohrad Basin, Ottnangian and Karpatian sediments; 7 – Upper Karpatian—Lower Badenian ryolite formation (Tarr Tuff);
8 – Middle to Upper Miocene volcanics; 9 – Middle to Upper Miocene sediments; 10 – Upper Miocene to Pleistocene basalts.
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Fig. 2. Lithostratigraphic scheme of the North Hungarian—South Slovakian basins, providing the correlation of the Oligocene—Lower Mio-
cene formations in the Lučenec, Ipe and Rimava Depressions (Vass & Elečko 1982 adapted by Holcová 2005) and Eger area (Báldi 1986).
Abbreviations: Mb – member, CGL – conglomerates, Lms. – limestones.
water sediments deposited in high-energy shoreface and shal-
low-marine environments described as the Čakanovce Beds,
Lipovany Sandstone, Tachty Sandstone, Jalová Sandstone and
Pétervására Sandstone (e.g. Vass & Elečko 1982; Sztanó &
Tari 1993). The Late Eggenburgian regression resulted in ter-
restrial clay and sandy-gravel deposition of the Zagyvapálfa
and Bukovinka Formations, which include numerous bird and
mammal footprints covered by thick tuff deposits of the
Gyulakeszi Rhyolite Tuff Formation (Márton et al. 1996).
The new subsidence is related to accommodation of the
Nógrád/Novohrad basin in the Early Ottnangian time. Ma-
rine transgression was preceded by fluvial, swampy and
lacustrine sedimentation of the Salgótarján Formation,
which contains coal bearing deposits of the Pôtor Member
(Vass et al. 1983). The upper Ottnangian—Karpatian forma-
tions of marine deposits are represented by the Plachtince/
Mátra Beds and Modrý Kameň/Egyházasgerge Formations.
The main part of the Modrý Kameň Formation belongs to the
Sečianky Member also named the Garáb Schlier, consisting
of calcareous claystones with full-marine fauna such as
benthic and planktonic foraminifers, mollusks, gastropods
(e.g. Holcová 1996; Ondrejíčková 1972; Vass 2002). The
Modrý Kameň Formation is superposed by the Tarr Tuff
Formation, which marks the beginning of volcanic activity at
the Karpatian-Badenian boundary. Badenian basin-fill forma-
tions are formed by shoreface sands of the Príbelce Member,
volcanoclastic sediments and volcanic intrusive complexes
named the Vinica Formation, Halič and Šiator Andesites and
Pokoradza Formation. The Nógrád/Novohrad Basin came to
an end due to domal uplift of the South Slovakian volcanic
area, which resulted in marine regression and basin inversion
from the Middle Badenian time (Vass & Šucha 1994; Vass
1995b; Kováč et al. 2001). Upper Miocene deposits of the
Lučenec and Rimava depressions are represented by fluvial
gravels and variegated clays of the Poltár Formation and ba-
salts of the Podrečany Formation.
Material and methods
The Rapovce GTL-2 core was drilled at a site located
4.36 km south of Lučenec near Rapovce village
(N 48°16’39.52”, E 19°40’43.47” – Fig. 1). The borehole
reached a depth of up to 1501.5 m and caught the following
lithostratigraphic formations (Vass et al. 2008 – Fig. 2):
Fi akovo Formation (5.00—45.00 m), Lučenec Formation
(45.00—606.00 m), Číž Formation (606.00—775.00 m) and
Mesozoic basement complex represented by the Middle- and
Upper Triassic carbonates (775.00—1501.50 m). The sands
and sandstones of the Fi akovo Formation in the upper part
of the GTL-2 borehole belong to the Lipovany Member. The
Lučenec Formation consists of grey calcareous siltstones of
the Széczény Schlier (45.00—595.00 m) with intercalations
of 1m thick beds of fine-grained sandstones. Sediments below
the Széczény Schlier belong to the Panica Member, consisting
of coarse- to medium grained conglomerates and sandstones
(595.00—606.00 m). The Číž Formation reaches a thickness of
up to 169 m (606.00—775.00 m) and its main part is formed by
deltaic sands of the Rapovce Member (606.00—745.00 m),
outer shelf siltstones and claystones of the Lenártovce Mem-
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ber (745.00—65.00 m), lagoon coal-bearing sandstones of the
Hostišovce Member (795.00—770.00 m) and transgressive
sandstones and pebbly sandstones of the Blh Member
(770.00—775.00 m).
Both the standard Mediterranean and regional Central
Paratethys stages to the Oligocene and Early Miocene strati-
graphic interpretation. The Paleogene Time Scale for the
standard Mediterranean stages followed Gradstein et al.
(2012) and the Paratethyan stages were applied in accor-
dance with their definition for the Kiscellian (Báldi 1969,
emend. Báldi et al. 1984), Egerian (Báldi & Seneš 1975,
emend. Báldi et al. 1999) and Eggenburgian (Steininger &
Seneš 1971). The Paratethyan nomenclature was also used in
an informal sense, as for designation of Kiscell-type fossil
assemblages, sedimentary facies, transgressive events (e.g.
initial flooding of the Kiscellian Sea into the South Slovakian
Basin – cf. Vass 2003).
The borehole section was sampled every 5 meters and pro-
vided 78 samples for foraminiferal and calcareous nanno-
plankton study. The samples were taken mainly from the
calcareous-rich claystones and siltstones. Nannofossil sam-
ples were prepared by the decantation method (Haq & Loh-
mann 1976; Perch-Nielsen 1985). The smear slides were
inspected under the light microscope Zeiss at 1.500
× magni-
fication and nannofossils documented by digital camera. For
quantitative analysis, at least 300 specimens per sample were
counted in randomly selected fields.
The foraminiferal microfauna was washed, dried and
sieved, obtaining the size fractions > 250 µm, > 150 µm and
> 38 µm. The residues were examined under a stereomicro-
scope Olympus and picked for study using the JEOL Scan-
ning Electron microscope.
Nannoplankton age determination was carried out accord-
ing to the zonation by Martini (1971), completed by Perch-
Nielsen (1985), Young (1998) and Wise et al. (2002).
Planktonic foraminiferal biostratigraphy was applied on the ba-
sis of the Oligocene—Early Miocene zonal schemes (Berggren
& Miller 1988; Spezzaferri 1994; Berggren et al. 1995) and
Paratethyan foraminiferal data (Sztrákos 1979; Rögl 1985;
Cicha et al. 1998; etc.). Bioevents of nannoplankton and fora-
miniferal biostratigraphy were denoted by the acronyms
sensu Berggren & Pearson (2005), “LO” for the lowest occur-
rence and “HO” for the highest occurrence of the index species.
determinable, G (good) – the species are very easily identi-
fiable, etching and overgrowth is slight.
The samples were evaluated by using the following abun-
dance scale: VH (very high) – more than 20 specimens in the
1 field of view, H (high) – 10—20 specimens in the 1 field of
view, M (moderate) – 5—10 specimens in the 1 field of view,
L (low) – 1—5 specimens in the 1 field of view and VL (very
low) – less than 5 specimens in the 5 field of view.
Climatic proxies and trophic conditions were inferred
from the temperature preferences and life strategy of calcare-
ous nannofossils (Wei et al. 1992; Bralower 2002; Persico &
Villa 2004; Bartol et al. 2008) and planktonic foraminifera
(Boersma & Premoli Silva 1983, 1991; Spezzaferi 1995;
Wade et al. 2007).
Results
Calcareous nannofossils
Preservation of calcareous nannofossils was moderate, be-
cause the specimens are slightly mechanically damaged, but
most of them can still be identified easily. Abundance of cal-
careous nannofossils was low, considering < 5 specimens oc-
cur in one field of view of the microscope.
Species diversity
Nannoplankton assemblages are moderately diversified,
involving a total number of 37 nannofossil taxa with
30 (85 %) autochthonous and 7 (15 %) reworked species.
Their proportion is rather equal, showing a slight downwar
increase of reworked species in the Číž Formation (18 %)
and upward increase of autochthonous species in the
Lučenec Formation (86 %) – Fig. 3. The ratio of the re-
worked species could have increased due to nannofosil re-
sedimentation in prodeltaic/deltaic sediments of the Číž
Formation, whereas the indigenous species dominated dur-
ing the stabilization of full-marine environments in the
Lučenec Formation (Széczény Schlier – cf. Vass 1995a).
Nannofossils of the Číž Formation from the depth
775—595 m are rich in species Coccolithus pelagicus, Cycli-
Fig. 3. Nannofossil variations in the ratio of autochthonous versus re-
worked species across the Rapovce GTL-2 section.
Nannofossil assemblages were evaluated on the basis
of specimen preservation and abundance, nannofossil
temperature preferences and their trophic strategy. Sta-
tistical methods (percentage analyses) were performed
by counting of 300 specimens on each sample, provid-
ing paleoecological data for evaluation of autochtho-
nous nannofossil species. Percentage share of each
nannoplankton group is expressed in diagrams.
Preservation of calcareous nannofossils was deter-
mined according to Roth & Thierstein (1972) as fol-
lows: VP (very poor) – etching and mechanical
damage to specimens is very intensive, P (poor) – frag-
mentation and/or overgrowth of the specimens makes it
difficult to identify them, M (moderate) – mechanical
damage and etching of the specimens are visible in the
minority of the nannoassemblage, specimens are easily
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cargolithus floridanus, Dictyococcites bisectus and Zygrha-
blithus bijugatus. The common species of this nannoplankton
association are Helicosphaera compacta, Helicosphaera
euphratis, Reticulofenestra lockeri and Reticulofenestra
scrippsae. There is also a low abundance of the species
Isthmolithus recurvus and Reticulofenestra ornata, which
disappear at the 595 m interval.
The most abundant nannofossils in the Lučenec Formation
comprise the species Coccolithus pelagicus, Cyclicargolithus
floridanus, Dictyococcites bisectus and Zygrhablithus biju-
gatus. They are commonly associated with Helicosphaera
compacta, Helicosphaera euphratis and Reticulofenestra
lockeri. The nannofossil association with Cyclicargolithus
abisectus, Helicosphaera recta and Pontosphaera desueta
appeared from the depth of 470—595 m. The association
changed at the depth of 470 m by disappearance of Heli-
cosphaera compacta and Sphenolithus distentus and appear-
ance of Pontosphaera enormis. Rare occurrences of Discoaster
cf. adamanteus, Triquetrorhabdulus cf. carinatus and Sphe-
nolithus conicus were recorded from the depth of 345—300 m.
Some important species of the association, such as Cyclicar-
golithus abisectus, Helicosphaera recta and Dictyococcites
bisectus, vanished at the depth of 210 m. New specimens of
nannofossils Helicosphaera carteri, Helicosphaera mediterra-
nea and Discoaster druggii appeared from the depth of 160 m.
Nannoplankton biostratigraphic interpretations
Nannofossil assemblages of the Číž and Lučenec forma-
tions consist of several stratigraphically important taxa,
which enable us to distinguish the following nannoplankton
zones (Figs. 4, 5):
The NP23 Zone was determined in the interval 775—595 m
based on the species Reticulofenestra ornata, R. lockeri and
Isthmolithus recurvus, which have their lowest common oc-
currence reported from this zone.
The appearance of Reticulofenestra lockeri is the typical
nannofossil datum of the NP23 Zone (Nagymarosy & Voronina
1992). This interval represents a peak of anoxic conditions
and stagnant regime in Paratethyan basins (Báldi 1984). The
isolation of Paratethys and the decline of salinity within the
NP23 Zone caused the extinction of cosmopolitan nannoflora
and expansion of the endemic species Reticulofenestra lockeri
that tolerated hyposaline waters (Nagymarosy 2000; Melinte-
Dobrinescu & Brustur 2008). The NP23/NP24 zone boundary
was stated on the basis of the first occurrence of Cyclicar-
golithus abisectus (Perch-Nielsen 1985; Young 1998). Some
authors reported this species already from the upper part of the
NP23 Zone (Fornaciari & Rio 1996; Kaenel & Villa 1996;
Maiorano & Monechi 2006; Melinte-Dobrinescu & Brustur
2008). In the borehole, the LO of C. abisectus was found at
595 m close to the boundary of the Číž and Lučenec forma-
tions (Fig. 5).
The NP24 Zone was delimited by the overlapping ranges
of stratigrafically important species Cyclicargolithus abisec-
tus, Helicosphaera recta, Pontosphaera desueta and Reticu-
lofenestra lockeri. The appearance of cosmopolitan species
reflects restored marine environments and sea-level rise
(Krhovský & Djurasinovič 1992; Švábenická & Stráník
2004). However, the nannoplankton diversity of the NP24
Zone is not markedly higher than that of the NP23 Zone.
The NP24—NP25 boundary is inferred from the HO of Sphe-
nolithus distentus (Berggren et al. 1995) and was recorded at
470 m. Pontosphaera enormis, which has its LO within
NP24—25, can be used as a marker species (Young 1998). In
the Rapovce section it has a depth of 470 m. This boundary
was also outlined by the species of Helicosphaera compacta,
which has occurrences at 470 m that imply their HO in the
NP24 Zone (Maiorano & Monechi 2006; Melinte-Dobrinescu
& Brustur 2008). The nannofossil associations of the NP24
and NP25 zones should also be different in size from Cycli-
cargolithus abisectus (Śliwińska et al. 2012), whereby the
abundance of Cyclicargolithus abisectus >10 µm corresponds
to the lower part of the NP24 and NP23 zones and large-sized
nannofossils of Cyclicargolithus abisectus >13 µm corre-
spond to the uppermost part of the NP24 and lower part of
NP25 zones. Significant increase in the size of these nanno-
fossils was not observed. The upper part of the NP 25 Zone is
marked by the LOs of Triquetrorhabdulus cf. carinatus and
Sphenolithus conicus in the interval between 345—300 m.
The Oligocene-Miocene boundary is unclear since there is
no agreement on the definition of the base of the NN1 Zone
(LO of Helicosphaera recta sensu Martini, 1971) and CN1
Zone (LO Sphenolithus ciperoensis sensu Okada & Bukry,
1980). Later, this boundary was redefined by the LO of Dic-
tyococcites bisectus at the base of the NN1 Zone (Berggren
et al. 1995; Fornaciari & Rio 1996) or by the FOs of Spheno-
lithus belemnos and Helicosphaera carteri in the upper part
of the NN1 Zone (Young 1998). These nannofossil datums
from the Mediterranean were completed by some additional
bioevents from the Paratethyan basins, where the Oligocene-
Miocene boundary can be marked by the HO of Dictyococ-
cites bisectus and FADs of Helicosphaera kampteri and
Helicosphaera scissura (Holcová 2001; Garecka 2012),
Reticulofenestra pseudoumbilica (Holcová 2005), Spheno-
lithus delphix (Rögl & Nagymarosy 2004), Helicosphaera
mediterranea (Marunteanu 1999; Chira 2004) and Spheno-
lithus capricornutus (Holcová 2005; Melinte-Dobrinescu &
Brustur 2008).
The Oligocene-Miocene boundary of the Rapovce section
was identified by the HOs of Helicosphaera recta and Dictyo-
coccites bisectus at the depth of 210 m. This datum is fol-
lowed by the appearance of Helicosphaeraceae from the NN1
and NN2 zones. The LOs of Helicosphaera carteri and Heli-
cosphaera mediterranea were recorded at the depth of 135 m.
The NN1—NN2 zone boundary is constrained by the LO of
Discoaster druggii dated to 23.2 Ma chronostratigraphic scale
(Berggren et al. 1995), and in Paratethyan stages corresponds
to the Egerian-Eggerburgian boundary (Lehotayová 1982;
Marunteanu 1999; Holcová 2002). The LO of Discoaster
druggii was recorded in the depth of 155 m. These bioevents
are too scattered to provide consistent indications of the Oli-
gocene-Miocene boundary (sensu Young 1998).
Nannofossil paleoecology
Temperature proxies: The average proportion of the
warm-water species, discoasterids, helicosphaerids and sphe-
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noliths forms 15 % with increased quantity in the interval of
775—595 m (21 %), corresponding to the NP23 Zone, Mid-
Rupelian. The next increase of the warm-water species (17 %)
was recorded in the interval 50—150 m, which is correlated with
the NN2 Zone, Eggenburgian. The group of species preferring
temperate-water conditions such as Cyclicargolithus florida-
nus, Cyclicargolithus abisectus, Coccolithus sp. and Dictyo-
coccites bisectus reaches 54 % of the assemblage (Fig. 6).
The percentage of the cold-water species, above other
reticulofenestrids is approximately 16 % of the nannofossil
assemblage. The proportion of these species varied by rhyth-
mical decreasing and inceasing with temperature fluctuation.
A slight decrease of the cold-water species to 13 % was ob-
served above 175 m.
The nannofossil distribution in the GTL-2 section suggests
water temperature differences between the Číž Formation
Fig. 4. Calcareous nannoplankton species from the Číž and Lučenec formations in the Rapovce GTL-2 section. 1 – Isthmolithus recurvus,
760 m, NP23 Zone, Číž Formation; 2 – Lanternithus minutus, 760 m, NP23 Zone, Číž Formation; 3 – Dictyococcites bisectus, 745 m,
NP23 Zone, Číž Formation; 4 – Reticulofenestra ornata, 745 m, NP23 Zone, Číž Formation; 5 – Cyclicargolithus floridanus, 725 m,
NP23 Zone, Číž Formation; 6, 7 – Cyclicargolithus abisectus, 6—560 m, 7—515 m, NP24—25 Zone, Lučenec Formation; 8 – Reticu-
lofenestra lockeri, 560 m, NP24—25 Zone, Lučenec Formation; 9 – Pontosphaera multipora, 515 m, NP24—25 Zone, Lučenec Formation;
10 – Pontosphaera desueta, 495 m, NP25 Zone, Lučenec Formation; 11, 12 – Pontosphaera enormis: 11 – 495 m, 12 – 375 m, NP24—25
Zone, Lučenec Formation; 13 – Helicosphaera euphratis, 545 m, NP24—25 Zone, Lučenec Formation; 14 – Helicosphaera mediterranea,
135 m, NN1—NN2 Zone, Lučenec Formation; 15 – Discoaster druggii, 155 m, NN2 Zone, Lučenec Formation, 16 – Helicosphaera carteri,
135 m, NN1—NN2 Zone, Lučenec Formation. Scale bars 1 µm.
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Fig. 5. Distribution of nannofossils in the Rapovce GTL-2 section with indications of important index species for biostratigraphic classifi-
cation of the Oligocene—Early Miocene formations.
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(Rupelian) and Lučenec Formation
(Chattian). In the Číž Formation, the
highest share of nannofossils belongs
to the temperate-water species (54 %),
complemented by relatively common
warm-water taxa (22 %) and a small
number of cold-water taxa (13 %). The
assemblages of the Lučenec Formation
were also formed mostly by temperate-
water species (54 %), but they were
complemented by few specimens of
warm-water taxa (13 %) and an in-
creased number of cold-water species
(17 %). Considering the above men-
tioned data, the sediments of the Číž
Formation were deposited in warmer
water conditions than the sediments of
the Lučenec Formation.
The balanced composition of the
nannofossil assemblages in the inter-
val from 535—770 m of the Lučenec
Depression indicates an increase of
warm water conditions during the Early
Oligocene. That is related to the onset
of the Kiscellian transgression (Soták
et al. 2002; Soták 2010). This period
corresponds to the NP23 and NP24
zones. Similar conditions were reported
from the Novaj section (Báldi-Beke
1980, 1984), the Harshegy Sandstone
Formation and the Kiscell Clay For-
mation (Báldi-Beke 1980). Early to
Middle Oligocene temperature varia-
tions were also recorded by nannofos-
sil proxies in the Krynica Zone of the
Magura Nappe (Oszczypko-Clowes &
Żydek 2012).
Samples from the interval between
50—150 m show an increase of warm-
water species related to Early Miocene
climate warming in the NN2 Zone
(Soták et al. 2002; Chira 2004; Švábe-
nická et al. 2007).
Trophic resources: Calcareous
nannofossils of the Rapovce section
imply
different
trophic-preference
conditions. The first group consists of
Fig. 6. Quantitative diagram of nannofossils
expressed as percentage of species with
warm-water preferences (A), temperate-
water preferences (B) and cold-water pref-
erences (C) plotted against the depth in the
Rapovce GTL-2 borehole section (nanno-
plankton temperature preferences according
to Wei et al. 1992; Bralower 2002; Persico
& Villa 2004; Bartol et al. 2008).
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Fig. 7. Quantitative diagram of nannofossils expressed as percentage of eutrophic species (A) and oligotrophic species (B) and plotted
against the depth in the Rapovce GTL-2 borehole section (nannoplankton trophic preferences according to Wei et al. 1992; Bralower 2002;
Persico & Villa 2004; Bartol et al. 2008).
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eutrophic species such as Braarudosphaera bigelowii, Cycli-
cargolithus floridanus, Cyclicargolithus abisectus, Dictyo-
coccites bisectus, Reticulofenestra ornata and Zyghrablithus
bijugatus (Aubry 1992; Krhovský et al. 1992; Villa et al.
2008).
The second group consists of oligotrophic species that are
able to survive even in the areas poorer in nutrients. Oligo-
trophic species are represented by autochthonous forms such
as Coccolithus, Discoaster, Helicosphaera, Reticulofenestra,
Sphenolithus, Thoracosphaera, Triquetrorhabdulus (Haq &
Lohman 1976; Wei & Wise 1990a,b; Aubry 1992; Bralower
2002).
The third group consists of species for which trophic strat-
egy is not exactly specified. Their quantity forms the missing
percentage in calculation of high- and low-nutrient taxa in
nannofossil assemblages.
Trophic preferences of calcareous nannofossils (Fig. 7)
show a slightly higher percentage of oligotrophic species
(44 %) over the high-nutrient taxa (42 %). Eutrophic species
are more present in the lower part of the section (48 %), where
they prevail over oligotrophic species (38 %). This interval
corresponds to the NP23 Zone. Contrary to this, the eutrophic
species are less frequent in the higher part of the borehole
(39 %), where the oligotrophic species become dominant
(49 %). Considering that, the nannofossils of the Číž Forma-
tion indicate more eutrophic conditions (48 %) than their
mostly oligotrophic assemblages in the Lučenec Formation
(39 %). The proliferation of eutrophic species could have been
caused by increased resources of nutrients in the basins during
the Kiscellian transgression (cf. Báldi-Beke 1980, 1984;
Švábenická et al. 2007). Similar eutrophic conditions have
also been identified in the Lower Oligocene formations of the
Magura basin (Oszczypko-Clowes & Żydek 2012).
Foraminifera: biostratigraphy and paleoenvironments
Planktonic foraminifera of the borehole section show a wide
species diversity and distinctive changes in frequency and dis-
tribution (Figs. 8, 9). Their succession reaches a great abun-
dance already at the base of the Číž Formation (770/750 m),
comprising large-sized forms of globigerinids, turborotaliids
and paragloborotaliids. Planktonic associations are diversified
to include the following species: Turborotalia ampliapertura,
Paragloborotalia nana, Subbotina gortanii, Globigerina
praebulloides, Globigerina officinalis and Globigerina leroi.
These foraminiferal species enable us to determine the Mid-
dle Rupelian age (O2 Zone sensu Berggren & Pearson
2005). Planktonic bioproductivity increased probably simul-
taneously with the sea-level rise during the Kiscellian trans-
gression. Transgressive conditions of the Číž Formation are
also indicated by well-aerated shelf environments of benthic
foraminifera, represented by the epifaunal species Eponides
umbonatus, Heterolepa dutemplei, Anomalina cryptomphala,
Lenticulina budensis, Percultazonaria fragaria. Microfau-
nistic assemblages of the Číž Formation changed in deltaic
sediments of the Rapovce Member in 620—750 m, where
planktonic foraminifers provide a distinct reduction in test-
size, abundance and diversity. Their association consists of
minute forms of tenuitellids, cassigerinellids, globorotaloids
and catapsydracids. A similar horizon of small-sized forms
of planktonic foraminifers was observed by Horváth (1998)
in the Hungarian Paleogene Basin. The taxonomic diversity
of planktonic foraminifers in the Rapovce Member decreases
to several species: Tenuitella gemma, Cassigerinella globu-
losa, Globorotaloides hexagonus and Catapsydrax martini.
Dwarfing of planktonic foraminfers implies stress-induced
conditions, such as the influence of hyposaline waters on the
morphogenesis of Cassigerinella chipolensis (Martinotti,
1989) or cool-water preferences of tunuitellids (Pipperr &
Reichenbacher, 2010). Stress conditions of surface-water
productivity, which were also indicated by the appearance of
pyritized frustules of diatoms and tests of pteropods, were
caused probably by eutrophication, shallowing and brackish-
ing of sea-water (Šutovská 1987). The decreased planktonic
productivity is replaced by increased benthic productivity,
which was manifested by the abundance of uvigerinid fora-
minifers. The population density of the Rapovce Member
was increased in some horizons by monospecific associa-
tions of Uvigerina hantkeni corresponding to “the Uvigerina
bloom” in the Kiscell Clay at the base of the NP24 Zone
(Horváth 1998).
The transitional beds between the Číž and Lučenec Forma-
tions from 600—460 m contain the Kiscellian-Egerian associ-
ations of foraminiferal microfauna. The character of the
microfauna implies the restoration of full-marine conditions,
which is manifested by blooming of the planktonic foramini-
fers and recolonization of the benthic biota. The abundance
of planktonic foraminifera increased abruptly at the begin-
ning of the transgressive cycle above the Rapovce Member
in 600—605 m, by the appearance of the Egerian species like
Globoturborotalita ouachitaensis, G. angulisuturalis, G.
woodi and G. labiacrassata (Holcová in Vass et al. 2004).
Those foraminifers are associated with Paragloborotalia
opima, P. bella, Globigerina praebulloides and Globigeri-
Fig. 8. Foraminiferal species from the Číž and Lučenec Formations in the Rapovce GTL-2 section. 1 – Turborotalia ampliapertura, 770 m,
O2 Zone, Číž Formation; 2 – Subbotina gortanii, 770 m, O2 Zone, Číž Formation; 3, 4 – Paragloborotalia opima, 515 m, O4—O5 Zone,
Číž Formation; 5, 6 – Globigerinoides primordius, 455 m, transitional beds of the Číž and Lučenec Formations, O5 Zone; 7, 8 – Globigerina
praebulloides: 7 – 55 m, M1 Zone, Lučenec Formation, 8 – 770 m, O2 Zone, Číž Formation; 9, 10 – Glogoturborotalita ouachitaensis,
605 m, O4 Zone, Číž Formation; 11 – Globoturborotalita woodi, 605 m, O4 Zone, Číž Formation; 12 – Globoturborotalita connecta,
455 m, O5 Zone, transitional beds of the Číž and Lučenec Formations; 13 – Paragloborotalia bella, 605 m, O4 Zone, Číž Formation;
14 – Paragloborotalia pseudokugleri, 455 m, O5 Zone, transitional beds of the Číž and Lučenec Formations; 15 – Globigerina wagneri,
455 m, O5 Zone, transitional beds of the Číž and Lučenec Formations; 16 – Globigerina anguliofficinalis, 225 m, O6 Zone, Lučenec For-
mation; 17, 18 – Globigerina ciperoensis, 135 m, O6 Zone, Lučenec Formation – upper part; 19, 20 – Globigerina ottnangiensis, 55 m,
M1 Zone, Lučenec Formation – uppermost part.
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Fig. 9. Distribution of foraminifers in the Rapovce GTL-2 section with indications of important index species for biostratigraphic classifi-
cation of the Oligocene—Early Miocene formations.
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nella obesa. Common occurrence of species from the Glo-
boturborotalita – group with G. angulisuturalis (LO in O4
Zone according to Berggren & Pearson 2005) and P. opima
indicates the Early Chattian biozones (P20—21/O3—O4 Zones
sensu Berggren & Miller 1988; Berggren & Pearson 2005).
Benthic foraminifers are represented by euryoxibiont and
low-oxygen tolerant species like Sphaeroidina bulloides,
Pulenia bulloides, Eggerella bradyi, Uvigerina vickburgen-
sis, Bulimina cf. altasica, Bulimina schischkinskayae, Cas-
sidulinoides tenuis, Kareriella quadrinoides, Stilostomella
consobrina and Stilostomella adolphina.
The base of the Szécsény Schlier of the Lučenec Formation
at 455 m is marked by the HO of Paragloborotalia opima.
Similarly, the presence of this species in the Lučenec Forma-
tion was also noted by Holcová (2001, 2005). The extinction
of P. opima provided an important datum for the early Chat-
tian biozone (O5 Zone sensu Berggren & Pearson 2005). At
the same horizon, the LO of Globigerinoides primordius was
recorded. This species appeared together with other primi-
tive specimens of the genus Globigerinoides such as transi-
tional forms to G. altiaperturus and G. immaturus (Keller
1981; Ouda 1998). Change to the new globigerinid forms
with secondary appertures on the spiral side appeared in re-
sponse to Late Oligocene warming (Jenkins 1973). The
planktonic foraminifers of the Lučenec Formation also in-
clude Globoturborotalita ouachitaensis, G. connecta, Globi-
gerina wagneri, Paragloborotalia pseudokugleri (LO in
O6 Zone with estimated age 25.9 Ma – Berggren et al.
1995), “Dentoglobigerina” cf. venezuelana and Globigeri-
nella obesa. The benthic foraminiferal microfauna of the
Lučenec Formation is enriched in epifaunal habitats with
plano-convex trochospiral, biconvex trochospiral, rounded
planispiral, spherical and uniserial morphotypes (sensu
Corliss & Chen 1988). They belong to the following species
Cibicidoides ungerianus, C. lopjanicus, Gyroidenoides sol-
dani, Eponides umbonatus, Fontbothia wuellerstorfi,
Sphaeroidina ciperana, Lenticulina cultrata, Percultazonaria
fragaria, Vaginulinopsis pseudodecorata, Anomalina affinis,
Ceratocancris haueri, Pullenia bulloides and Stilostomella
pauperata. The predominance of epifaunal and shallow infau-
nal benthic foraminifers implies improved conditions of bot-
tom water oxygenation in the the Lučenec Formation.
The foraminiferal microfauna changes in the upper part of
the Lučenec Formation from 330—320 m, with calcareous
species diminishing and agglutinated species becoming
dominant. Associations involve species like Trochammina
squamata, T. inflata, T. globigeriniformis and Haplophrag-
moides sp. This assemblage suggests the deep-water associa-
tion of agglutinated foraminifera with a reduced amount of
dissolved oxygen. Deep-water conditions of the middle
bathyal zone are also indicated by infaunal benthic species
like Bulimina coprolithoides, Pullenia bulloides and Globo-
cassidulina depressa (cf. Kaiho & Nishimura 1992). These
species also provide evidence of dysoxic and highly produc-
tive conditions (Corliss & Chen 1988). The occurrence of
trochamminid species and bathyal benthic foraminifers in the
upper part of the Lučenec Formation indicates a deepening-
upward basin during the Egerian time. This is also evident
from a high concentration of siliceous sponge rhaxes in
schlier sediments. Sea-level rise in the Lučenec Formation
coincides with the global eustatic cycle TB1.4, which culmi-
nated in the topmost part of the Széczeny Schlier with the
appearance of late Oligocene to early Miocene microfauna
(Šutovská-Holcová et al. 1993; Vass 1995a).
Late Oligocene planktonic foraminifera were recovered
with the appearance of Paragloborotalia pseudokugleri at
225 m. It extended to the topmost part of the P22/O6 Zone
(Spezzaferri 1991). This species is associated with Globigerina
ciperoensis, G. anguliofficinalis, Globigerinoides primordius
and G. praealtiaperturus, their common occurrences are
known from the O6 Zone. This evidence appears to establish
the Late Chattian age of the Lučenec Formation at 225 m.
The globigerinid foraminifers in the uppermost part of the
Lučenec Formation (above 225 m) are developed mostly as
five-chambered species belonging to Globigerina ottnang-
iensis. The appearance of this species can be used as a bio-
stratigraphic marker of the Oligocene-Miocene boundary
(Rögl 1994; Cicha et al. 1998). The abundance maximum of
G. ottnangiensis occurs at 55 m. Early Miocene associations
of planktonic foraminifera are completed by Cassigerinella
globulosa, Paragloborotalia semivera, “Dentoglobigerina”
venezuelana and Globigerina praebulloides. Benthic associ-
ations are formed mostly by epifaunal forms with rounded
planispiral, plano-convex and biconvex trochospiral, and
spherical morphotypes. They belong to the species: Angulo-
gerina angulosa, Lenticulina arcuatostriata, L. cultrata,
Cibicidoides ungerianus, C. lopjanicus, Almaena osnabru-
gensis, Hanzawia boueana, Melonis affinis, and Pullenia
bulloides. Infaunal species are less frequent, for example
Praeglobobulimina pupoides and Globocassidulina oblonga,
but some of them already belong to the Early Miocene fora-
minifera. The Eggenburgian index species Uvigerina post-
hantkeni (cf. Cicha et al. 1998; Rupp & Haunold-Jenke
2003; Grunert et al. 2013) appeared in the uppermost part of
the Lučenec Formation at 55 m.
Conclusions
The Oligocene and Early Miocene deposits of the Číž and
Lučenec Formations provide important data for recognition
of the planktonic bioevents and paleoenvironments of the
South Slovakian Basin (Fig. 10). The data from the Rapovce
GTL-2 section lead to the following conclusions:
1. The Kiscellian transgression at the beginning of the
NP23 Zone is marked by extinction and speciation of nanno-
fossils (e.g. LO R. lockeri and R. ornata and HO I. recurvus),
and diversification and high-rate bioproductivity of plank-
tonic foraminifera (paragloborotaliids, turborotaliids, globi-
gerinids). The transgressive cycle corresponds to the TB1.2
event of global eustasy (Vass 1995a), which is documented
by well-aerated shelf environments with epifaunal benthic
foraminifers and nannofossil proxies of warm- to temperate-
water and oligotrophic conditions.
2. A regressive phase in the Late Rupelian is documented
by the lowstand deposition of prodeltaic sediments of the
Rapovce Member. These sediments reveal stress-induced
conditions due to eutrophication and brackishing of sea-wa-
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ter, which resulted in diversity fall and blooming of endemic
nannoflora, especially Reticulofenestra ornata, and dwarfing
of planktonic foraminifera such as tenuitellids and catapsy-
dracids. Infaunal biota, mainly uvigerinid foraminifers, pro-
liferated from increased nutrients and bottom-water dysoxia.
Improvement of the open-marine conditions occurred within
the NP23—24 boundary, which is approximated by the ap-
pearance of nannofossil species Cyclicargolithus abisectus.
3. Restoration of full-marine conditions was controlled by
the sea-level rise of the TB1.3 cycle in the transitional se-
quences of the Číž and Lučenec formations. Transgressive
sediments are enriched with reworked nannofossils (Fig. 3).
The autochthonous species of the NP24 Zone include cosmo-
politan nannofossils such as Cyclicargolithus, Helicosphaera,
Dictyococcites and Pontosphaera (Fig. 6). The climatic prox-
Fig. 10. Integrated biostratigraphy of the Rapovce section with datum events of planktonic foraminifers (black triangles) and nannofossils
(white triangles). Biostratigraphic data are indicated by the highest occurrences (HO) and lowest occurrences (LO) of the index taxa.
ies of nannofossils indicate eutrophic and temperate-water
conditions. Planktonic foraminiferal assemblages with Para-
globorotalia opima and Globigerina ciperoensis define the
O5 Zone. The abundance of globoturborotaliid and para-
globorotaliid species, which belong to mixed-layered or upper
thermocline habitats (Pearson & Wade 2009), implies warm-
to temperate-water water conditions. Oxygen depleted con-
ditions are indicated by deep-infaunal benthic species like
Bulimina, Chilostomella and Stilostomella.
4. The Rupelian-Chattian boundary was identified by the
HO of Paragloborotalia opima and the HO of Sphenolithus
distentus at the base of the Lučenec Formation (O5 Zone,
NP24—25 Zone) (Fig. 10). New taxa of planktonic foraminifer
are represented by globigerinid forms with secondary aper-
tures (the LO of Globigerinoides primordius), which are asso-
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ciated with five-chambered species (Globigerina ciperoensis)
and the first paragloborotaliid forms with arched sutures (P.
pseudokugleri). Both the appearance of primitive Globigeri-
noides and the nannofossil Sphenolithus distentus provide
evidence of the Late Oligocene climatic warming. Nannofos-
sil assemblages of the Lučenec Formation are also enriched
in content of oligotrophic and temperate-water species com-
pared to those in the Late Rupelian sediments of the Číž For-
mation. The benthic microfauna of the Lučenec Formation
consists of mostly epifaunal and in the upper part also agglu-
tinated forms, which provide evidence of better oxygenation
and deepening-upward conditions during the sea-level rising
of the TB1.4 cycle.
5. The Oligocene-Miocene boundary was identified in the
uppermost part of the Lučenec Formation at 210 m, by sev-
eral nannofossil bioevents such as the HOs of Helicosphaera
recta and Dictyococcites bisectus and LOs of Helicosphaera
carteri, H. mediterranea and Discoaster druggii in the NN1
and NN2 zones (Fig. 10). Early Miocene planktonic fora-
minifera are represented mostly by five-chambered species
belonging to Globigerina ottnangiensis. Benthic microfauna
strongly increased in abundance with large-sized tests of
Lenticulina species, associated with numerous specimens of
Angulogerina angulosa and Uvigerina posthantkeni. Benthic
life could proliferate with opening of the Eggenburgian sea-
way connection into the Lučenec Depression. This became
evident mainly in the Fi akovo/Pétervására Basin (Sztanó
1995; Kováč et al. 2001).
Acknowledgment: The authors are grateful to S. Ćorić and
M. Oszczypko-Clowes for their constructive comments and
improvements of the manuscript. Our thanks go to D. Vass
and J. Dzúrik for providing the opportunity to study the
Rapovce borehole section and their advice and assistance.
This work has been supported by the research agency VEGA
Project 2/0042/12, and by funds received through the Centre
of Excellence for Integrative Research of the Earth’s Geo-
sphere (ITMS 262201200064).
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Checklist of taxa mentioned in the text in alphabetic order:
Appendix 1
Nannoplankton species
Arkhangelskiella cymbiformis Vekshina, 1959
Braarudosphaera bigelowii (Gran & Braarud, 1935) Deflandre,
1947
Coccolithus eopelagicus (Bramlette & Riedel, 1954) Bramlette
& Sullivan, 1961
Coccolithus formosus (Kamptner, 1963) Wise, 1973
Coccolithus pelagicus (Wallich, 1877) Schiller, 1930
Coronocyclus nitescens (Kamptner, 1963) Bramlette &
Wilcoxon, 1967
Cyclicargolithus abisectus (Müller, 1970) Wise, 1973
Cyclicargolithus floridanus (Hay et al., 1967) Bukry, 1971
Dictyococcites bisectus (Hay, Mohler & Wade, 1966) Bukry
& Percival, 1971
Discoaster adamanteus Bramlette & Wilcoxon, 1967
Discoaster deflandrei Bramlette & Riedel, 1954
Discoaster druggii Bramlette & Wilcoxon, 1967
Helicosphaera ampliaperta Bramlette & Wilcoxon, 1967
Helicosphaera bramlettei (Müller, 1970) Jafar & Martini, 1975
Helicosphaera carteri (Wallich, 1877) Kamptner, 1954
Helicosphaera compacta Bramlette & Wilcoxon, 1967
Helicosphaera euphratis Haq, 1966
Helicosphaera kampteri (Hay & Mohler in Hay et al., 1967),
Locker, 1970
Helicosphaera mediterranea Müller, 1981
Helicosphaera recta (Haq, 1966) Jafar & Martini, 1975
Helicosphaera scissura Miller, 1981
Holodiscolithus macroporus (Deflandre in Deflandre & Fert,
1954) Roth, 1970
Isthmolithus recurvus Deflandre in Deflandre & Fert, 1954
Pontosphaera desueta (Müller, 1970) Perch-Nielsen, 1984
Pontosphaera enormis (Locker, 1967) Perch-Nielsen, 1984
Pontosphaera multipora (Kamptner, 1948) Roth, 1970
emend. Burns, 1973
Reticulofenestra bisecta (Hay, Mohler & Wade, 1966) Roth,
1970
Reticulofenestra lockeri Müller, 1970
Reticulofenestra minutula (Gartner, 1967) Haq & Berggren,
1978
Reticulofenestra ornata Müller, 1970
Reticulofenestra pseudoumbilica (Gartner, 1967) Gartner, 1969
Reticulofenestra scrippsae (Bukry & Percival, 1971) Roth,
1973
Reticulofenestra umbilica (Levin, 1965) Martini & Ritz-
kowski, 1968
Sphenolithus belemnos Bramlette & Wilcoxon, 1967
Sphenolithus capricornutus Bukry & Percival, 1971
Sphenolithus ciperoensis Bramlette & Wilcoxon, 1967
Sphenolithus conicus Bukry, 1971
Sphenolithus delphix Bukry, 1973
Sphenolithus distentus (Martini, 1965) Bramlette & Wilcoxon,
1967
Sphenolithus moriformis (Brönnimann & Stradner, 1960)
Bramlette & Wilcoxon, 1967
Transversopontis pygmaea (Locker, 1967) Perch-Nielsen, 1984
Zygrhablithus bijugatus (Deflandre in Deflandre & Fert,
1954) Deflandre, 1959
Appendix 2
Foraminiferal species
Almaena osnabrugensis (Roemer, 1838)
Angulogerina angulosa (Willimson, 1858)
Anomalina affinis (Hantken, 1948)
Anomalina cryptomphala (Reuss, 1850)
Bulimina cf. Altasica Cushman & Parker, 1937
Bulimina coprolithoides Andreae, 1884
Bulimina schischkinskayae Samoylova, 1947
Cassidulinoides tenuis Phleger & Parker, 1951
Cassigerinella globulosa (Egger, 1857)
Cassigerinella chipolensis (Cushman & Ponton, 1932)
Catapsydrax martini (Blow & Banner, 1962)
Ceratocancris haueri (d’Orbigny, 1839)
Cibicidoides lopjanicus (Myatlyuk, 1950)
Cibicidoides ungerianus (d’Orbigny, 1846)
“Dentoglobigerina” venezuelana (Hedberg, 1937)
Eggerella bradyi (Cushman, 1911)
Eponides umbonatus (Reuss, 1860)
Fontbothia wuellerstorfi (Schwager, 1866)
Globigerina anguliofficinalis (Blow, 1969)
Globigerina ciperoensis Bolli, 1954
Globigerinella obesa (Bolli, 1957)
Globigerina ottnangiensis Rögl, 1969
Globigerina officinalis Subbotina, 1953
Globigerina praebulloides Blow, 1959
Globigerina praebulloides leroi Blow & Banner, 1962
Globigerina wagneri Rögl, 1994
Globigerinoides altiaperturus Bolli, 1957
Globigerinoides immaturus Le Roy, 1939
Globigerinoides primordius Blow & Banner, 1962
Globocassidulina depressa (Asano & Nakamura, 1937)
Globocassidulina oblonga (Reuss, 1850)
Globorotaloides hexagonus Natland, 1938
Globoturborotalita angulisuturalis (Bolli, 1957)
Globoturborotalita brazieri (Jenkins, 1966)
Globoturborotalita connecta (Jenkins, 1964)
Globoturborotalita labiacrassata (Jenkins, 1966)
Globoturborotalita ouachitaensis (Howe & Wallace,
1932)
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EOLOGICA CARPA
OGICA CARPA
OGICA CARPA
OGICA CARPA
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THICA
THICA
THICA
THICA, 2014, 65, 6, 451—470
Globoturborotalita woodi Jenkins, 1960
Gyroidenoides soldani d’Orbigny, 1982
Hanzawia boueana (d’Orbigny, 1846)
Heterolepa dutemplei (d’Orbigny, 1846)
Kareriella guadryinoides (Fornasini, 1885)
Lenticulina arcuatostriata (Hantken, 1875)
Lenticulina budensis (Hantken, 1875)
Lenticulina cultrata (Montfort, 1808)
Melonis affinis (Reuss, 1851)
Paragloborotalia bella (Jenkins, 1967)
Paragloborotalia nana (Bolli, 1957)
Paragloborotalia opima (Bolli, 1957)
Paragloborotalia pseudocontinuosa (Blow, 1959)
Paragloborotalia pseudokugleri (Blow, 1969)
Percultazonaria fragaria (Gümbel, 1868)
Praeglobobulimina pupoides (d’Orbigny, 1846)
Pulenia bulloides(d’Orbigny, 1826)
Sphaeroidina bulloides (d’Orbigny, 1846)
Sphaeroidina ciperana Cushman & Todd, 1949
Stilostomella adolphina (d’Orbigny, 1846)
Stilostomella consobrina (d’Orbigny, 1839)
Stilostomella pauperata (d’Orbigny, 1839)
Subbotina gortanii (Borsetti, 1959)
Tenuitella gemma Jenkins, 1966
Trochammina globigeriniformis (Parker & Jones, 1865)
Trochammina inflata (Montagu, 1808)
Trochammina squamata (Jones & Parker, 1860)
Turborotalia ampliapertura Bolli, 1957
Uvigerina hantkeni (Cushman & Edwards, 1937)
Uvigerina posthantkeni Papp, 1971
Uvigerina vickburgensis (Cushman & Ellisor, 1931)
Vaginulinopsis pseudodecorata Hagn & Holzl, 1952