GeolCarp_Vol59_No4_319

GEOLOGICA CARPATHICA, AUGUST 2008, 59, 4, 319—332

www.geologicacarpathica.sk

Introduction

The term “Podhale Paleogene” refers to a part of the Central
Carpathian  Paleogene  deposits  (Fig. 1;  see  also  Soták  et  al.
2001) that occur in Poland between the Pieniny Klippen Belt
in the north and the Tatra Mountains in the south (Fig. 2). The
northern  margin  of  the  Podhale  Paleogene  along  the  contact
with  the  Pieniny  Klippen  Belt  is  tectonic  (e.g.  Birkenmajer
1960,  1970,  1985)  whereas  its  southern  margin  is  erosional.

Middle-Late Eocene phytoplankton from marl intraclasts

(Podhale Paleogene, Inner Carpathians, Poland):

biostratigraphic and paleoenvironmental implications

PRZEMYSŁAW GEDL

and MAŁGORZATA GARECKA

Institute of Geological Sciences, Cracow Research Centre, Polish Academy of Sciences, Senacka 1, 31-002 Kraków, Poland;

ndgedl@cyf-kr.edu.pl

Polish Geological Institute, Carpathian Branch, Skrzatów 1, 31-560 Kraków, Poland; malgorzata.garecka@pgi.gov.pl

(Manuscript received July 11, 2007; accepted in revised form February 13, 2008)

Abstract: Organic-walled dinoflagellate cysts and calcareous nannoplankton are described from marly intraclasts found in
submarine slump deposits within the Lower Oligocene Szaflary Beds exposed in the Leśnica Stream. Their Middle and
Late Eocene age implies that the investigated deposits are coeval with the basal deposits of the Podhale Paleogene succes-
sion. These Middle and Upper Eocene marl intraclasts were eroded and transported into the flysch basin during the Early
Oligocene. They represent the sediments deposited in the northern part of the Podhale Basin that is not exposed in recent
times. Paleoenvironmental analysis of microfossils suggests sea-level oscillations during late Middle-Late Eocene (Bartonian—
Priabonian) with its maximum during the earliest Late Eocene (earliest Priabonian). A drop of sea surface temperature
during Late Eocene is also suggested on the basis of high-latitude microfossil occurrence.

Key words: Eocene, Carpathians, Poland, Central Carpathian Paleogene, Podhale Basin, paleogeography, biostratigraphy,
calcareous nannoplankton, dinoflagellate cysts.

As  a  consequence,  the  present-day  northernmost  peripheries
of the Podhale Basin are now unknown because they are tec-
tonically disturbed along the boundary with the Pieniny Klip-
pen  Belt  structure  (e.g.  Birkenmajer  1970,  1985).  The
present-day  northernmost  deposits  of  the  Podhale  Paleogene
that crop out along the contact with the Pieniny Klippen Belt
are  the  Lower  Oligocene  Szaflary  Beds  –  the  Eocene  rocks
are exposed in contact with the Pieniny Klippen Belt deposits
only in Slovakia (Orava and Haligovce area; see e.g. Gross &

Fig. 1. Tectonic sketch-map of the northern part of the Central Carpathian Paleogene Basin with location of the study area (arrowed).

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GEDL and GARECKA

Köhler 1987; Birkenmajer 1979 respectively). The basal strata
of the Podhale Paleogene in this area, represented by the so-
called  Tatra  Eocene  (presumably  Middle-Upper  Eocene)  are
only  known  from  boreholes  (e.g.  Kępińska  1997).  However,
incomplete coring and poor microfossil content of these basal
intervals  makes  the  recognition  of  this  interval  limited.  Dis-
covery of coeval marl intraclasts found in a submarine slump
layer within the Szaflary Beds (Gedl 2004a) brings new infor-
mation  on  the  age,  paleoenvironment  and  paleogeographical
position  of  the  basal  deposits  of  the  Podhale  Paleogene.  For
this  purpose  dinoflagellate  cysts  and  calcareous  nannoplank-
ton from marly intraclasts were studied.

Geological setting

The Podhale Paleogene sequence is traditionally divided

into two informal units: the so-called Tatra Eocene (or Num-
mulitic Eocene) in the lower part (Borové Formation sensu
Gross et al. 1984) and the so-called Podhale Flysch in the

upper part. The basal part of the Tatra Eocene is usually de-
veloped  as  conglomerates  of  delta  and  cliff  breccias  origin
(e.g. Passendorfer 1958) passing upwards into variously de-
veloped  carbonates  (e.g.  Roniewicz  1969).  Marly  deposits
that occur locally in the topmost part of the Tatra Eocene se-
quence  (Alexandrowicz  &  Geroch  1963;  Bartholdy  et  al.
1999) are more frequent in the northern part of the Podhale
Basin  (Sokołowski  1992;  Jaromin  et  al.  1992;  Kępińska
1997). The thickness of the Tatra Eocene rarely exceeds 100
meters (exceptional maximal thickness above 300 meters oc-
curs in the Hruby Regiel Mountain area).

The Podhale Flysch deposits, resting upon the Tatra

Eocene  or  locally  directly  on  the  Mesozoic  substrate,  reach
up  to  3000—3500 m  thickness.  This  unit  was  informally  di-
vided by Gołąb (1959) and Watycha (1959) into the Szaflary
Beds occurring in the northern part of the Podhale Basin, the
Zakopane  Beds  (Huty  Formation  sensu  Gross  et  al.  1984),
the Chochołów Beds (Zuberec Formation sensu Gross et al.
1984) and the Ostrysz Beds known from the eastern part of
the Podhale Basin (Biely Potok Formation sensu Gross et al.

Fig. 2. Location of the study area (arrowed) against the background of a simplified sketch-map of the Podhale Paleogene (after Małecka
1982; with correction in Tatra Mts by Birkenmajer 2000).

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MIDDLE-LATE EOCENE PHYTOPLANKTON (PODHALE PALEOGENE, INNER CARPATHIANS, POLAND)

1984). The Szaflary Beds, the oldest Podhale Flysch deposits
in the peri-Klippen area, are developed as proximal flysch
sediments. They are covered by the Zakopane Beds, the old-
est flysch deposits in the peri-Tatric area.

The Middle-early Late Eocene (Bartonian—early Priabon-

ian) age of the Tatra Eocene carbonates was determined on
the  basis  of  large  and  small  foraminifers  (e.g.  Bieda  1963;
Olszewska & Wieczorek 1998). The higher part of the Tatra
Eocene exposed in Pod Capkami quarry was dated by means
of  calcareous  nannoplankton  as  NP16—NP17  (Middle
Eocene:  upper  part  of  Lutetian—Bartonian;  Bartholdy  et  al.
1995). Some uncertainties occur concerning the dating of the
uppermost part of the Tatra Eocene where frequent plankton-
ic foraminifers were found (Alexandrowicz & Geroch 1963;
Blaicher 1973; Olszewska & Wieczorek 1998; Bartholdy et
al. 1999).

The age of the Szaflary Beds, often synonymized with the

Šambron Beds sensu Chmelík (1957), based on dinocysts is
Early  Oligocene  (but  not  the  earliest  Oligocene;  Fig. 3).  A
younger,  but  also  Early  Oligocene  age  was  determined  for
the Zakopane Beds (Gedl 1999, 2000a). As a consequence, a
hiatus  embracing  the  uppermost  Eocene  to  lower  part  of
Lower  Oligocene  was  suggested  between  the  Tatra  Eocene

and Podhale Flysch deposits. The hiatus is wider in the south-
ern  part  of  the  Podhale  Basin,  where  it  also  includes  time
equivalents  of  the  Szaflary  Beds  (Gedl  2000a).  A  similar
“mid-Oligocene” age of the Zakopane Beds was concluded by
Garecka (2005) who correlated this lithostratigraphic unit with
the NP24 Zone. The younger Chochołów Beds are correlated
with  the  upper  part  of  the  Lower  Oligocene  (Rupelian;  di-
noflagellate cysts; Gedl 2000a) and Upper Oligocene (Chatti-
an; foraminifers and calcareous nannoplankton; Olszewska &
Wieczorek 1998; Garecka 2005 respectively).

Material

The slump layer with investigated marly intraclasts is ex-

posed on the left bank of the Leśnica Stream (Fig. 4; see also
Gedl  2004a).  It  occurs  within  thin-bedded  flysch  deposits
with a distinctive bentonite layer a few meters above it. Few
other  slump  layers  are  exposed  in  its  vicinity  (see  Gedl
2000a, p. 81). The investigated slump layer is composed of a
1.2-m  thick  exotic-bearing,  conglomeratic  layer  passing
sharply  upwards  into  1.5-m  thick  sandstone  layer  (Fig. 5).
The conglomeratic layer consists of well-rounded clasts (up

Fig. 3. Age of Podhale Paleogene (dinoflagellate cysts after Gedl 2000a; calcareous nannoplankton after Garecka 2005).

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GEDL and GARECKA

to  several  centimeters  in  diameter)  representing  mainly  the
Tatra-derived  Triassic  dolomites.  Among  those  hard  peb-
bles,  infrequent  soft  cream  yellow  and  pale  brown,  up  to
10 cm in diameter, marly clasts of irregular shape occur. Six
samples were taken from these marly clasts (LśnE1—LśnE6).
Additionally, two samples were taken from the surrounding
Szaflary  Beds  shales:  sample  LśnŁ1  from  just  below  and
sample LśnŁ2 from just above the slump layer (Fig. 5).

Methods

Dinoflagellate cysts. The samples for palynological inves-

tigation were processed following the standard palynological
procedure including 38% hydrochloric acid (HCl) treatment,
40% hydrofluoric acid (HF) treatment, heavy liquid
(ZnCl

+ HCl; density 2.0 g/cm

) separation, ultrasound for

10—15 s and sieving at 15 µm nylon-mesh. Two slides were
made from each sample using glycerine jelly as a mounting
medium. All dinoflagellate cysts from both slides were de-
termined and counted. Photographs were taken with the us-
ing a Sony DSC-S75 camera.

Samples, palynological residues and slides are stored in

the collection of the Institute of Geological Sciences, Polish
Academy of Sciences, Kraków.

Calcareous nannoplankton. Samples were prepared as

smear slides. Calcareous nannoplankton was investigated
with the light microscope at a magnification of 1500

×, and

photographed using a H-III Nikon camera.

Results

Dinoflagellate cysts (Figs. 6A, 7). Palynofacies of the

flysch shales from just below and above the slump layer
(Szaflary Beds; samples LśnŁ1 and LśnŁ2) is composed of
land plant tissue remains and sporomorphs (mainly bisaccate
pollen grains; a similar palynofacies dominated by terrestrial
elements is characteristic for this part of the Szaflary Beds:
see Gedl 2000a). Single specimens of Deflandrea phosphor-
itica and Wetzeliella sp. represent the extremely rare di-
noflagellate cysts.

Marly intraclasts from the slump layer are characterized by

different palynofacies. Terrestrial phytoclasts are represented
by  small-sized  opaque  phytoclasts  and  structured  tissue  re-
mains.  A  different  palynofacies  than  in  other  marly  clasts
was found in sample LśnE4 taken from a soft brownish marl.
It  is  composed  predominately  of  hyalinous  resin  particles.
Sporomorphs and plant tissue remains are subordinate.

Dinoflagellate cysts are frequent, except in sample LśnE4,

and are, often the dominating palynomorph. They also differ
qualitatively being dominated by chorate taxa (Fig. 7). The
taxonomic characteristics of the dinocyst assemblage can be
found in Gedl (2004a).

Calcareous nannoplankton (Figs. 6B, 8). No calcareous

nannoplankton  (sample  LśnŁ1)  or  single,  very  poorly  pre-
served  specimens  of  Dictyococcites  bisectus  were  found
(sample  LśnŁ2)  in  the  Szaflary  Beds.  This  contrasts  with
much richer, although also poorly preserved, calcareous nan-

Fig. 4. Location of exposure of the investigated submarine slump
layer (after Gedl 2004a).

Fig. 5. Lithology of the investigated Szaflary Beds section with
submarine slump layer (after Gedl 2004a), including position of in-
vestigated samples.

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GEDL and GARECKA

noplankton assemblages from the marly clasts (samples
LśnE1 to LśnE6, except LśnE4).

A very rich calcareous nannoplankton assemblage domi-

nated  by  Dictyococcites  bisectus,  Discoaster  barbadiensis,
Discoaster  saipanensis,  Reticulofenestra  umbilica  and
Zygrhablithus  bijugatus  is  found  in  sample  LśnE1.  In  the
same sample Coccolithus pelagicus, Corannulus germanicus
and  Coccolithus  formosus  occur  frequently.  The  state  of
preservation  is  very  poor.  Damaged,  broken  specimens  are
the  most  common;  their  precise  determination  is  often  im-
possible.  This  especially  refers  to  the  representatives  of  the
cold-water genus Chiasmolithus. Broken remains of large spe-
cies  of  this  genus,  Chiasmolithus  cf.  grandis  and  Chiasmo-
lithus  cf.  oamaurensis,  are  found  in  sample  LśnE1  and  are
the only determinable remains of this genus. Representatives
of Prinsiaceae, Dictyococcites bisectus and Reticulofenestra
umbilica, are also found as broken remains only. Numerous
representatives  of  Prinsiaceae  are  known  to  be  resistant  to
dissolution. They often occur together with  Coccolithus pe-
lagicus as the only taxa in the impoverished Tertiary calcare-
ous  nannoplankton  assemblages.  Corranulus  germanicus
also shows the traces of dissolution. Representative of Heli-
cosphaeraceae, Helicosphaera compacta is also very poorly
preserved but in this case, there was no difficulty with deter-
mination of this species, although they are mainly preserved
as the central parts only. Warm-water Discoasteraceae show
damage  to  arms,  one  of  their  diagnostic  features.  There  are
some  very  small  representatives  of  Sphenolithaceae  ob-
served  in  this  sample.  Unfortunately,  their  small  size  and
poor  state  of  preservation  made  suprageneric  determination
impossible.

Sample LśnE2 yielded an impoverished calcareous nanno-

plankton  assemblage.  Infrequent  specimens  of  Coccolithus
formosus,  Coccolithus  pelagicus  and  Zygrhablithus  bijuga-
tus are found. Reticulofenestra dictyoda, Reticulofenestra cf.
umbilica,  Sphenolithus  aff.  furcatolithoides  and  Spheno-
lithus spiniger occur as single specimens only.

More frequent nanofossils are found in sample LśnE3.

Calcareous nannoplankton from this sample shows very poor
preservation,  mostly  mechanic  breaking.  This  refers  mainly
to  the  Prisniaceae  (i.e.  Reticu-
lofenestra and Dictyococcites) as
well  to  the  Coccolithaceae  and
Discoasteraceae.  The  latter  are
infrequent  and  often  have  dam-
aged  or  missing  arms.  Remains
of large representatives of Retic-
ulofenestra,  most  likely  Reticu-
lofenestra umbilica, are found in
this sample.

Sample LśnE4, was the only

one from the marly clast samples,
to  yield  no  calcareous  nanno-
plankton.  Two  further  samples,
LśnE5  and  LśnE6,  contain  rich,
although poorly preserved calcar-
eous  nannoplankton  assemblage.
Assemblage  of  sample  LśnE5
contains  Discoaster  tanii,  Dis-

coaster cf. tanii nodifer and Helicosphaera cf. compacta. The
one from sample LśnE6 is dominated by Dictyococcites bisec-
tus,  Reticulofenestra  umbilica,  Discoaster  barbadiensis, Dis-
coaster saipanensis, Discoaster tanii, Zygrhablithus bijugatus
and Chiasmolithus sp. All specimens are poorly or very poor-
ly  preserved.  The  latter  especially  refers  to  the  very  large
forms of Reticulofenestra umbilica. The genus Chiasmolithus
is  also  represented  by  very  large  forms  preserved  as  incom-
plete specimens, often difficult to determine – Chiasmolithus
aff.  grandis  and  Chiasmolithus  aff.  oamaruensis.  Numerous
isolated arms of discoasters and fragments of multiarm forms
of  Discoaster  barbadiensis  and  Discoaster  saipanensis  are
found in this sample.

Interpretation

Age of microfloral assemblages

The age of the dinocyst assemblages (Fig. 9) is based on

comparison with dinocyst stratigraphic distribution in noth-
western Europe (e.g. Bujak & Mudge 1994) and the Mediter-
ranean (e.g. Brinkhuis & Biffi 1993) as well comparison with
Paleogene dinocyst distribution in Middle-Upper Eocene and
Oligocene strata of the Polish Carpathians (Bujak in Van Cou-
vering 1981; Gedl 1996, 2000a,b, 2004a,b,c, 2005).

The first attempts to introduce of calcareous nannoplank-

ton  zonations  of  Tertiary  deposits  were  undertaken  in  the
early  sixties  (Brönnimann  &  Stradner  1960;  Bramlette  &
Sullivan 1961). Several other zonal schemes were proposed
a  few  years  later,  for  example  by  Radomski  (1967,  1968),
Martini  (1971)  and  Bukry  (1973,  1975;  the  latter  modified
later by Okada & Bukry 1980). Two of them, those of Marti-
ni  (1971)  and  Okada  &  Bukry  (1980),  became  widely  ac-
cepted  and  are  now  in  common  use.  Their  usefulness  is
related  to  the  widespread  occurrences  of  the  zonal  marker
taxa  making  possible  a  correlation  among  widely  separated
areas.  The  zonation  scheme  proposed  by  Okada  &  Bukry
(1980) is more useful in low latitude areas since most of the
index taxa are tropical forms. Martini’s zonation is, in turn,

Fig. 9. Comparison of dinocyst- and calcareous nannoplankton-age interpretation of the studied
marly clasts.

327

MIDDLE-LATE EOCENE PHYTOPLANKTON (PODHALE PALEOGENE, INNER CARPATHIANS, POLAND)

more helpful in higher latitudes. The latter scheme is there-
fore used in the present study (Fig. 9).

The inferred age of the calcareous nannoplankton assem-

blage from sample LśnE1 is the earliest Late Eocene. The as-
semblage is characteristic for the NP18 Zone (Chiasmolithus
oamaruensis  Zone)  of  Martini  (1971).  This  is  based  on  the
presence  of  Chiasmolithus  cf.  oamaruensis,  the  lowest  oc-
currence  of  which  defines  the  base  of  the  NP18  Zone,  and
Corannulus  germanicus  and  Orthozygus  aureus  that  have
their lowest occurrence within the NP18 Zone. Isthmolithus
recurvus, a diagnostic taxon for the NP19—20 Zones (sensu
Martini  1976),  has  not  been  found  in  this  sample.  Several
taxa found in sample LśnE1 have their lowest occurrences in
the NP17 Zone (uppermost Middle Eocene): Discoaster tanii
(lowest  occurrence  in  the  middle  part  of  this  zone),  Heli-
cosphaera  compacta  and  Sphenolithus  predistentus.  The
presence  of  Clathrolithus  aff.  ellipticus  and  Clathrolithus
aff.  spinosus  was  noted.  These  species  are  believed  to  be
characteristic  for  the  latest  Middle  Eocene  (Aubry  1988),
whereas Perch-Nielsen (1985) reports Clathrolithus spinosus
as a Late Eocene species. A slightly younger age can be con-
cluded for sample LśnE1 on the basis of its dinocyst assem-
blage.  Lack  of  Areosphaeridium  michoudii  in  this  sample,
and  the  presence  of  Areosphaeridium  diktyoplokum  imply
Late  Eocene  (middle  or  late  Priabonian)  age.  However,  in
the light of calcareous nannoplankton interpretation, the ab-
sence of Areosphaeridium michoudii could be accidental.

The age of sample LśnE2 cannot be precisely estimated. It

contains very infrequent calcareous nannoplankton and
poorly preserved dinoflagellate cysts among which no diag-
nostic species were identified. Reticulofenestra cf. umbilica,
the youngest species identified in this sample, has its lowest
occurrence within the NP16 Zone. This suggests a latest
Middle Eocene age for this assemblage.

The presence of Areosphaeridium michoudii in sample

LśnE3 indicates that this sample is no younger than NP19—20.
In  the  same  sample  Dracodinium  laszczynskii  occurs.  This
species was described from the Middle Eocene and the lower
part of Upper Eocene strata in the Flysch Carpathians (Gedl
1996, 2005). A similar age can be concluded for this sample
on  the  basis  of  the  presence  of  Melitasphaeridium  pseu-
dorecurvatum  that  occur  in  this  sample  only.  This  species
has  the  highest  occurrence  in  the  early  Late  Eocene  (top  of
the NP18 Zone; Stover et al. 1996). Presence of Discoaster
cf. tanii in sample LśnE3 suggests the latest Middle Eocene
age of this sample. The lowest occurrence of this species is
found  in  the  middle  part  of  the  NP17  Zone.  The  lower
boundary of this zone is defined by the highest occurrence of
Chiasmolithus solitus, its upper limit is based on the lowest
occurrence  of  Chiasmolithus  oamaruensis.  The  NP17  Zone
is correlated with Discoaster saipanensis Subzone of Okada
& Bukry (1980). Its lower boundary is defined by the highest
occurrences  of  Chiasmolithus  solitus  and  Discoaster  bifax,
its  upper  boundary  is  defined  by  the  highest  occurrence  of
Chiasmolithus grandis and the lowest occurrence of Chiasmo-
lithus oamaruensis. None of these index taxa were found in
the sample LśnE3. Only the presence of Discoaster cf. tanii
allows attribution to the middle part of the NP17 Zone. The
dinoflagellate  cysts  of  sample  Lśn3  resemble  dinocyst  as-

semblages found in deposits overlying the Pucov Conglom-
erate at Pucov (Slovakia) and correlated with NP18—20
Zones (Soták et al. 2007).

The age of sample LśnE4 remains uncertain. This sample

contains  no  calcareous  nannoplankton,  and  very  rare  di-
noflagellate cysts. This is presumably due to restricted envi-
ronmental  conditions.  The  dinocyst  species  present  in  this
sample  (Wetzeliella  symmetrica  and  Enneadocysta  pectini-
formis)  are  long  ranging.  Thus,  an  upper  Middle  Eocene
(Bartonian)—Oligocene age-assessment of this sample can be
concluded.

A latest Middle Eocene age (NP17 Zone), similar as that

of the sample LśnE3, is accepted for the sample LśnE5. This
is  based  on  the  presence  of  Discoaster  tanii,  Discoaster  cf.
tanii  nodifer  and  Helicosphaera  cf.  compacta,  species  that
have  their  lowest  occurrences  in  the  NP17  Zone.  As  in  the
sample LśnE3, no index species of older and younger zones
(i.e.  Chiasmolithus  solitus  and  Chiasmolithus  oamaruensis)
were found. In sample LśnE5 a presence of Sphenolithus cf.
pseudoradians was noted. The lowest occurrence of this spe-
cies,  according  to  Martini’s  zonation,  defines  the  upper
boundary of the NP19 Zone (i.e. the lower boundary of the
following NP20 Zone; the uppermost Upper Eocene). How-
ever, stratigraphic value of this species as an index taxon is
limited.  This  species  was  also  noted  from  older,  Middle
Eocene, deposits (NP15 Zone), and it is often difficult to dis-
tinguish  from  older  Sphenolithus  radians,  especially  in  the
case  of  poorly  preserved  material.  The  dinoflagellate  cysts
found in this sample cannot a precise age. Only the presence
of Aiora sp. A, which was also found in sample LśnE1 might
indicate the same age as the latter sample. This would imply
a slightly younger age (NP18).

The presence of Areosphaeridium diktyoplokum and the

lack of Areosphaeridium michoudii in sample LśnE6 suggest
the latest Eocene age of this assemblage. The highest occur-
rence of Areosphaeridium diktyoplokum is a widely accepted
marker of the Eocene-Oligocene boundary (e.g. Williams et
al. 1993; Stover et al. 1996). This implies that the age of the
sample  cannot  be  younger  than  Eocene.  The  highest  occur-
rence  of  Areosphaeridium  michoudii  in  the  North  Sea  de-
fines the top of the E8a Subzone of Bujak & Mudge (1994)
correlated  with  the  top  of  the  NP18  Zone.  However,  the
highest occurrence of Areosphaeridium michoudii in the Pol-
ish Carpathians is found in the NP19—20 Zone (Gedl 2005).
This, in turn, delimits the lower age boundary of the sample
LśnE5 that cannot be older than the NP19—20 Zone. Age in-
terpretation  of  calcareous  nannoplankton  from  this  sample
implies  the  latest  Eocene  age  (NP19—20  Zone).  This  inter-
pretation  is  based  on  the  co-occurrence  of  Isthmolithus  re-
curvus, Discoaster barbadiensis and Discoaster saipanensis.
The  lowest  occurrence  of  Isthmolithus  recurvus  defines  the
lower  boundary  of  the  NP19—20  Zone  (Aubry  1983).  The
same event defines the lower boundary of the NP19 Zone in
the scheme of Martini (1971). After fusion of the NP19 and
NP20 Zones, Martini (1976) used the highest occurrence of
Chiasmolithus  grandis  as  the  event  that  defines  the  lower
boundary  of  the  fused  NP19—20  Zone.  The  upper  boundary
of this zone is defined as the highest occurrence of Discoast-
er saipanensis (Martini 1971; Aubry 1983), a taxon present

328

GEDL and GARECKA

in the sample LśnE6. The presence of Cribrocentrum reticu-
latum was noted in this sample. The highest occurrence of
this species is known in lower latitudes from the top of the
NP18 Zone, whereas it is reported from Oligocene strata in
the higher latitudes. Thus, the end of the latest Eocene (late
Priabonian) age is correlated with the higher part of the
NP19—20 Zone.

The above-presented marl clast age-assessment indicates

that they are recycled. Although the age of the surrounding
flysch shales (samples LśnŁ1 and LśnŁ2) could not be pre-
cisely determined during this study due to lack of diagnostic
species, Gedl (2000b) accepted a Early Oligocene age of the
Szaflary Beds on the basis of the occurrence of species such
as Chiropteridium galea, Chiropteridium lobospinosum and
Wetzeliella gochtii in neighbouring outcrops.

Paleoenvironment

Dinocyst and calcareous nannoplankton assemblages

found  in  the  marl  intraclasts  have  been  studied  for  recon-
struction  of  sedimentary  conditions  during  the  Middle-Late
Eocene in this part of the Central Carpathian Paleogene Ba-
sin.  Interpretation  of  the  environmental  preferences  of  Ter-
tiary  and  recent  dinoflagellate  cysts  is  based  on  works  by
several authors, including Brinkhuis (1994), Dale (1996) and
Rochon  et  al.  (1999).  That  of  calcareous  nannoplankton  is
based mainly on studies of Báldi-Beke (1984), Wei & Wise
(1990), Aubry (1992) and Nagymarosy & Voronina (1992).
Three  aspects  of  paleoenvironment  are  discussed  here:  dis-
tance from the shoreline, salinity and climatic fluctuations.

The oldest sample LśnE2 contains a dinocyst assemblage

that seems to have inhabited shelf waters. There are no ocean-
ic dinoflagellate cysts. The presence of near shore Homotryb-
lium  tenuispinosum  is  rather  a  result  of  hydrodynamic
transport,  which  is  also  implied  from  palynofacies  that  con-
tains frequent terrestrial elements. The composition of calcare-
ous  nannoplankton  generally  confirms  the  offshore
depositional  setting.  Coccolithus  pelagicus,  Reticulofenestra
dictyoda and Sphenolithus sp., all found in this sample, are be-
lieved  to  be  offshore  taxa.  Their  climatic  interpretation  is
somewhat confusing. Coccolithus pelagicus is interpreted as a
cold-water  species,  Reticulofenestra  dictyoda  as  temperate-
water, and Sphenolithus sp. is a warm-water genus. However,
this assemblage may be interpreted rather as cold- to temper-
ate-water because there are no representatives of Discoaster, a
warm-water genus that occurs frequently in younger samples.

Different phytoplankton assemblages characterize two

samples representing the NP17 Zone: LśnE3 and LśnE5. The
samples: LśnE5 contains a dinocyst assemblage qualitatively
similar  to  that  from  sample  LśnE2.  The  most  frequent  are
chorate taxa representing genera Spiniferites and Operculod-
inium,  whereas  no  Impagidinium  was  found.  Additionally,
Hystrichokolpoma spp. and Achomosphaera alcicornu occur
in  this  sample.  The  calcareous  nannoplankton  assemblage
differs significantly by frequent occurrence of the warm-wa-
ter genus Discoaster, which is characteristic of open waters.
Helicosphaera  cf.  compacta,  another  warm-water  taxon
found in this sample, is rather a near shore species, possibly
transported into a more remote basin part. Further expansion

of the basin is recorded in phytoplankton assemblages from
sample  LśnE3.  Oceanic  dinoflagellate  cysts  (Impagidinium
spp., Corrudinium incompositum) occur for the first time in
the  investigated  material.  Most  of  the  calcareous  nannofos-
sils represent offshore taxa (e.g. Coccolithus pelagicus, Dis-
coaster sp., Sphenolithus sp.). Moreover, the palynofacies of
this  sample  is  dominated  by  dinoflagellate  cysts  implying
pelagic sedimentation. The presence of Pontosphaera scissu-
ra  and  Braarudosphaera  bigelowii,  and  Wetzelielloideae
among the dinoflagellate cysts, all believed to be near-shore
taxa associated with reduced salinity, suggests a low-salinity
environment in the peripheral part of the basin. Warm-water
conditions are suggested by the presence of Discoaster sp.

Similar paleogeography must have characterized this part

of  the  Podhale  Basin  also  during  the  earliest  Late  Eocene
(early Priabonian; NP 18 Zone; sample LśnE1). An offshore
environment  is  indicated  by  the  presence  of  the  oceanic  di-
noflagellate  cysts  Impagidinium  spp.  and  Nematosphaerop-
sis lemniscata, and offshore nannofossils Discoaster sp. and
Coccolithus  pelagicus.  Near-shore  taxa  (e.g.  Helicosphaera
compacta, Homotryblium spp.), some of them characteristic
of  reduced  salinity  environments  (e.g.  Wetzelielloideae,
Braarudosphaera  bigelowii,  Neococcolithes  minutus),  were
transported into the distal part of the basin. The occurrence
of cold-water Chiasmolithus may indicate a drop in tempera-
ture of surface waters during the early Priabonian, although
warm-water  nannofossils  like  Discoaster  have  also  been
found in this assemblage.

The dinocyst assemblage from the youngest sample LśnE6

contains  frequent  chorate  gonyaulacoids  but  it  differs  from
the  dinocyst  assemblages  in  the  older  samples  by  relatively
common occurrence of peridinioids (Deflandrea spp. and the
Wetzelielloideae). Oceanic dinoflagellate cysts are very rare
(single Impagidinium specimen). Frequent occurrence of pe-
ridinioids  during  the  latest  Late  Eocene  (late  Priabonian)
might be related to changes in nutrient availability in the sur-
face waters of the Podhale Basin. This may be related to the
reduced salinity in the near shore waters shown, for example,
by Braarudosphaera bigelowii and Neococcolithes minutus.
The  extension  of  the  basin  could  be  the  same  as  during  the
earliest Late Eocene (early Priabonian) – calcareous nanno-
plankton  assemblage  is  dominated  by  offshore  taxa  (Dis-
coaster  sp.,  Sphenolithus  sp.,  Reticulofenestra  sp.).
Occurrence  of  Isthmolithus  recurvus,  another  cold-water
species,  may  indicate  a  further  drop  of  temperature  of  sur-
face  waters  in  the  Podhale  Basin  during  the  latest  Late
Eocene (late Priabonian).

A different environment is indicated for the marly deposits

represented by sample LśnE4. Its palynofacies dominated by
terrestrial elements, mainly the hyalinous resins suggests a
very near shore depositional setting. Because the age of this
sample has not been precisely determined its relation to the
other samples is not certain.

Discussion

Marly intraclasts investigated in this paper represent the

remains of upper Middle—Upper Eocene (Bartonian—Pria-

329

MIDDLE-LATE EOCENE PHYTOPLANKTON (PODHALE PALEOGENE, INNER CARPATHIANS, POLAND)

bonian) deposits from northern, no longer existing part of the
Podhale Basin that became tectonically damaged during for-
mation of the Pieniny Klippen Belt structure (see Birkenma-
jer  1985).  The  paleogeographical  location  of  the  original
sediments in the northern part of the basin is based on analy-
sis of paleocurrent directions in the Leśnica Stream (Krysiak
1976).  Occurrence  of  Eocene  intraclasts,  together  with  re-
mains of Mesozoic substrate in submarine slump deposit, in-
dicates  extensive  erosion  of  the  northern  border  of  the
Podhale Flysch Basin during the Early Oligocene. This pre-
sumably took place on the tectonically controlled submarine
ramp  that  was  concluded  as  the  depositional  model  for  the
Szaflary Beds by Wieczorek (1989).

In Poland, the basal deposits of the present-day northern

part of the Podhale Basin were drilled by the Biały Dunajec
PAN-1  and  Bańska  IG-1  boreholes  (Jaromin  et  al.  1992;
Sokołowski  1992;  Kępińska  1997).  Large  and  small  fora-
minifers found in the Tatra Eocene from Bańska IG-1 were
interpreted  as  Middle  Eocene  (Kulka  1983;  Olszewska  &
Wieczorek  1998).  No  microfossils  were  found  in  the  Biały
Dunajec  PAN-1  succession  except  for  damaged  shells  of
nummulites (Jaromin et al. 1992). The Tatra Eocene deposits
are developed here as conglomerates (Biały Dunajec PAN-1;
Jaromin et al. 1992) or conglomerates passing upwards into
pelitic  limestones  (Bańska  IG-1;  M.  Cieszkowski  in  J.
Sokołowski 1992). However, due to poor core recovery, es-
pecially in the Bańska IG-1 borehole, the complete lithologi-
cal development is not certainly known. Marly deposits that
occur locally in the topmost part of the Tatra Eocene succes-
sion  are  hard  and  rather  dark  coloured  (Alexandrowicz  &
Geroch  1963;  Sokołowski  1973;  Olszewska  &  Wieczorek
1998) so they significantly differ in lithology from the soft,
pale  beige  marly  intraclasts  from  the  Leśnica  Stream.  Thus
the  investigated  marls  represent  a  unique  lithofacies  within
the Podhale Paleogene succession in Poland. The only litho-
logically  comparable  deposits  are  the  cream  yellow  marls
found  by  S.  Sokołowski  (in  Blaicher  1973,  p. 120)  at  the
base of the peri-Tatric Zakopane Beds exposed in the  Przy-
porniak Stream (see also Gedl 2000a, p. 98, fig. 23). Blaich-
er (1973) compared foraminifers from these sediments with

Fig. 10. Age of the investigated marl intraclasts and conceptual sea surface level and temperature changes.

the ones from the Upper Eocene Globigerina Marl of the
Outer Carpathians. Olszewska & Wieczorek (1998) dated
the marly deposits as foraminiferal Zones P15—P16 (i.e. up-
permost Middle—Upper Eocene) that generally agree with the
results of the present paper.

Changes of dinocyst and calcareous nannoplankton assem-

blages  and  palynofacies  most  likely  reflect  the  sea  surface
changes  in  the  Central  Carpathian  Paleogene  Basin  during
the Middle and Late Eocene (Fig. 10). The most offshore pa-
leoenvironment recorded in NP18 Zone (possibly “the high-
stand”  phase)  is  preceded  and  followed  by  more  inshore
settings  recorded  in  NP16?—NP17  and  NP19—20  Zones  re-
spectively (possibly fall of the sea surface). This agrees well
with the correlation of the eustatic curve with the lithostrati-
graphic  division  of  the  Central  Carpathian  Paleogene  Basin
deposits shown by Soták et al. (2001, fig. 10). The paleocli-
matic interpretations of these authors also fit well the scenar-
io  presented  in  this  paper.  Late  Middle  Eocene  (Bartonian)
phytoplankton  assemblages  were  replaced  by  cold-water
ones during the Late Eocene (Priabonian).

Comparison of described in this paper phytoplankton as-

semblages  with  the  ones  from  other  localities  may  lead  to
some regional paleogeographical conclusions. The similarity
in  taxonomical  diversity  of  dinocyst  assemblages  described
from Outer Carpathians coeval deposits (Gedl 2004c, 2005)
suggests  the  existence  of  sea-way  connections  between  the
Inner  and  Outer  Carpathian  basins  during  the  Middle  and
Late Eocene. Analysis of younger, Oligocene phytoplankton
assemblages  from  the  Podhale  Flysch  deposits  (Gedl
2000a,b;  Garecka  2005)  shows  significant  differences  with
the ones from the Oligocene of the Outer Carpathians. Both
dinocyst  and  calcareous  nannoplankton  assemblages  from
the  Oligocene  Menilite-Krosno  Beds  are  extremely  taxo-
nomically  impoverished  or  even  absent  (e.g.  Gedl  1999,
2004c; Garecka 2005; Gedl & Leszczyński 2005). This sug-
gests various paleoenvironmental conditions in those basins
during  the  Oligocene  and  possibly  the  Early  Miocene.  A
possible reason might have been a closure of the connections
between  the  Outer  and  Inner  Carpathian  basins  during  the
earliest Oligocene.

330

GEDL and GARECKA

Conclusions

1. Soft, pale beige marly intraclasts found within a subma-

rine  slump  layer  within  the  Oligocene  Szaflary  Beds  repre-
sent  Middle-Upper  Eocene  deposits.  Their  presence  here
indicates  erosion  of  the  northern  part  of  the  Podhale  Basin
substrate  during  the  Early  Oligocene.  These  marls  are  the
only known remain of the basal parts of the Podhale Paleo-
gene succession in this part of the Central Carpathian Paleo-
gene Basin.

2. The marly intraclasts contain generally rich dinocyst

and calcareous nannoplankton assemblages. Interpretation of
their ages (latest Middle-Late Eocene) agrees with dating of
the basal intervals of the Podhale Paleogene succession from
more southern parts of the basin.

3. Paleoenvironmental interpretation suggests that the

most offshore sedimentary setting recorded during the earli-
est Late Eocene (early Priabonian; NP18 Zone) was preced-
ed and followed by more inshore paleoenvironments. A
relative drop of the sea surface temperature during the Late
Eocene (Priabonian; NP18—20 Zones) is suggested in rela-
tion to warmer surface waters during the late Middle Eocene
(Bartonian; NP17 Zone).

Acknowledgments:  We  would  like  to  thank  Lilian  Švábe-
nická,  Aida  Andreyeva-Grigorovich,  and  an  anonymous  re-
viewer  for  reading  the  manuscript;  their  critical  remarks
significantly improved the manuscript.

References

Alexandrowicz S. & Geroch S. 1963: Association de petits Fora-

minif res dans l’Eoc ne de la Tatra. Ann. Soc. Géol. Pol. 33,
219—228 (in Polish with French summary).

Aubry M.P. 1983: Corrélations biostratigraphiques entre les forma-

tions paléog nes épicontinentales de l’Europe du Nord-Ouest,
basées sur la nannoplancton calcaire. Th se Université Pierre
et Marie Curie, Paris 6, 208, 83—08.

Aubry M.P. 1988: Handbook of Cenozoic calcareous nannoplank-

ton. Book 2: Ortholithae (Catinasters, Ceratoliths, Rhabdo-
liths). Micropaleontology Press, Amer. Mus. Natur. Hist., New
York, 1—279.

Aubry M.P. 1992: Late Paleogene calcareous nannoplankton evolu-

tion: a tale of climatic deterioration. In: Prothero D.R. & Berg-
gren W.A. (Eds.): Eocene—Oligocene climatic and biotic
evolution. Princeton University Press, Princeton, New Jersey,
272—309.

Bartholdy J., Bellas S.M., Ćosović V., Fuček V.P. & Keupp H.

1999:  Processes  controlling  Eocene  mid-latitude  larger  Fora-
minifera  accumulations:  modelling  of  the  stratigraphic  archi-
tecture  of  a  fore-arc  basin  (Podhale  Basin,  Poland).  Geol.
Carpathica 50, 435—448.

Bartholdy J., Bellas S.M., Mertmann D., Machaniec E. & Manutso-

glu E. 1995: Fazies-Entwicklung und Biostratigraphie einer
Sequenz eozäner Sedimente im Steinbruch Pod Capkami,
Tatra-Gebirge, Polen. Berliner Geowiss. Abh. E16, 409—425.

Báldi-Beke M. 1984: The nannoplankton of the Transdanubian Palaeo-

gene Formations. Geol. Hung., Ser. Palaeont. 43, 3—307.

Bieda F. 1963: Larger foraminifers of the Tatra Eocene. Inst. Geol.,

Prace 37, 1—215 (in Polish with English summary).

Birkenmajer K. 1960: Geological map of the Pieniny Klippen Belt,

1 : 10,000, Sheet Niedzica. Instytut Geologiczny, Warszawa
(explanations in Polish).

Birkenmajer K. 1970: Geological map of the Pieniny Klippen Belt,

1 : 10,000, sheet Bór na Czerwonem-Szaflary. Instytut Geolo-
giczny, Warszawa (explanations in Polish).

Birkenmajer K. 1979: Geological guide in the Pieniny Klippen Belt.

Wydawnictwa Geologiczne, Warszawa, 1—236 (in Polish).

Birkenmajer K. 1985: Fourth day. In: Birkenmajer K. (Ed.): Main

geotraverse  of  the  Polish  Carpathians  (Cracow-Zakopane).
Guide  to  Excursion  2.  Carpatho-Balkan  Geological  Associa-
tion,  13th  Congress,  Cracow,  Poland  1985.  Geol.  Inst.,
Warszawa, 90—136.

Birkenmajer K. 2000: Gosau-type conglomerate in the Rusinowa

Polana area, Polish Tatra Mts: its relation to the Lower Subtat-
ric Nappe. Bull. Polish Acad. Sci., Earth Sci. 48, 117—133.

Blaicher J. 1973: Microfauna of the Podhale Flysch in the Zakopane

IG 1 borehole. Biul. Inst. Geol. 265, 105—133 (in Polish with
English summary).

Bramlette M.N. & Sullivan F.R. 1961: Coccolithophorids and relat-

ed nannoplankton of the early Tertiary in California.
Micropaleontology 7, 129—188.

Brinkhuis H. 1994: Late Eocene to Early Oligocene dinoflagellate

cysts from the Priabonian type-area (northeast Italy); bios-
tratigraphy and palaeoenvironmental interpretation. Palaeo-
geogr. Palaeoclimatol. Palaeoecol. 107, 121—163.

Brinkhuis H. & Biffi U. 1993: Dinoflagellate cyst stratigraphy of

the Eocene/Oligocene transition in central Italy. Mar. Micro-
paleont. 22, 131—183.

Brönnimann P. & Stradner H. 1960: Die Foraminiferen- und Dis-

coasteridenzonen von Kuba und ihre interkontinentale Korrela-
tion. Erdöl-Zeitschrift 76, 364—369.

Bujak J. & Mudge D. 1994: A high-resolution North Sea Eocene di-

nocyst zonation. J. Geol. Soc., London 151, 449—462

Bukry D. 1973: Low-latitude coccolith biostratigraphic Drilling

Project. Initial Reports 15, 685—703.

Bukry D. 1975: Coccolith and silicoflagellate stratigraphy, north-

western Pacific Ocean, Deep Sea Drilling Project Leg 32. In:
Larson R.L., Moberly R. et al. (Eds.): Deep Sea Drilling
Project. Initial Reports 32, 677—701.

Chmelík F. 1957: Note sur l’étude géologique du Paléog ne des Car-

pates centrales dans la région de Šariš entre Šambron et Sabinov.
Zprávy o geologických výzkumech 1957, 81—88 (in Czech).

Dale B. 1996: Dinoflagellate cyst ecology: modelling and geologi-

cal applications. In: Jansonius J. & McGregor D.C. (Eds.): Pa-
lynology: principles and applications. Amer. Assoc. Stratigr.
Palynol. Found., Dallas, Texas, 3, 1249—1275.

Garecka M. 2005: Calcareous nannoplankton from the Podhale Fly-

sch (Oligocene-Miocene, Inner Carpathians, Poland). Stud.
Geol. Pol. 124, 353—369.

Gedl P. 1996: Middle Eocene dinoflagellate cysts from the

Rogoźnik section, Flysch Carpathians, Poland. Acta Palaeo-
bot. 35, 195—231.

Gedl P. 1999: The age of base and top of the Podhale Palaeogene

Flysch (Inner Carpathians, Poland) based on dinocysts. Bull.
Pol. Acad. Sci., Earth Sci. 47, 77—102.

Gedl P. 2000a: Biostratigraphy and palaeoenvironment of the

Podhale Palaeogene (Inner Carpathians, Poland) in the light of
palynological studies. Part I. Stud. Geol. Pol. 117, 69—154.

Gedl P. 2000b: Biostratigraphy and palaeoenvironment of the

Podhale Palaeogene (Inner Carpathians, Poland) in the light of
palynological studies. Part II. Summary and systematic de-
scriptions. Stud. Geol. Pol. 117, 155—303.

Gedl P. 2004a: Eocene dinoflagellate cysts from exotic clasts in

submarine slump, Podhale Flysch (Oligocene), Inner Car-
pathians, Poland. Stud. Geol. Pol. 123, 199—222.

331

MIDDLE-LATE EOCENE PHYTOPLANKTON (PODHALE PALEOGENE, INNER CARPATHIANS, POLAND)

Gedl P. 2004b: Dinoflagellate cysts from Šambron beds surround-

ing the Pucov Member in Orava, Central Carpathian Palaeo-
gene, Slovakia. Stud. Geol. Pol. 123, 223—243.

Gedl P. 2004c: Dinoflagellate cyst record of the Eocene-Oligocene

boundary succession in flysch deposits at Leluchów, Car-
pathian Mountains, Poland. In: Beaudoin A.B. & Head M.J.
(Eds.): The palynology and micropalaeontology of boundaries.
Geol. Soc. London, Spec. Publ. 230, 309—324.

Gedl P. 2005: Late Eocene-early Oligocene organic-walled di-

noflagellate cysts from Folusz, Magura Nappe, Polish Car-
pathians. Acta Palaeobot. 45, 27—83.

Gedl P. & Leszczyński S. 2005: Palynology of the Eocene-Oligocene

transition in the marginal zone of the Magura Nappe at Folusz
(Western Carpathians, Poland). Geol. Carpathica 56, 155—167.

Gołąb J. 1959: On the geology of the western Podhale Flysch area.

Biul. Inst. Geol. 149, 223—239 (in Polish with English summary).

Gross P. & Köhler E. 1987: Eocene transgression in Orava part of

the Pieniny Klippen Belt. Geol. Práce, Spr. 86, 157—164 (in
Slovak).

Gross P., Köhler E. & Samuel O. 1984: New lithostratigraphic divi-

sion of the Inner Carpathian Palaeogene. Geol. Práce, Spr. 81,
103—117.

Jaromin A., Kępińska B., Nagel J., Sokołowski J. & Wieczorek J.

1992: Geosynoptic documentation of the geothermal well
Biały Dunajec PAN-1. Geosynoptyka i Geotermia 2, 1—121 (in
Polish).

Kępińska B. 1997: Geologic-geothermal model of the Podhale area.

Centrum Podstawowych Problemów Gospodarki Surowcami
Mineralnymi i Energią PAN, Studia, Rozprawy, Monografie,
48, 1—111 (in Polish).

Krysiak Z. 1976: Directions of the transports of material in the

Podhale Flysch on the basis of data from the basin of the
Leśnica Stream (Eastern Podhale). Kwart. Geol. 20, 323—330
(in Polish with English summary).

Kulka A. 1983: Large foraminifera from the Tatra Eocene of the

Bańska IG-1 well. Kwart. Geol. 27, 871—872 (in Polish).

Małecka D. 1982: Map of main geological units of Podhale and sur-

rounding areas (1 : 100,000). Wydawnictwa Geologiczne,
Warszawa (in Polish).

Martini E. 1971: Standard Tertiary and Quaternary calcareous nan-

noplankton zonation. In: Farinacci A. (Ed.): Proceeding of the
Planktic Conference, Roma, 1970. Edizioni Tecnoscienza,
Roma, 1971, 739—785.

Martini E. 1976: Cretaceous to Recent calcareous nannoplankton

from the Central Pacific Ocean (DSDP Leg 33). Deep Sea
Drilling Project, Initial Reports 33, 383—423.

Nagymarosy A. & Voronina A.A. 1992: Calcareous nannoplankton

from the Lower Maykopian Beds (Early Oligocene, Union of In-
dependent States). Knihovnička ZPN 14b, 189—221.

Okada H. & Bukry D. 1980: Supplementary modification and intro-

duction of code numbers to the low-latitude coccolith biostrati-
graphic zonation (Bukry 1973, 1975). Mar. Micropaleontology
5, 321—325.

Olszewska B.W. & Wieczorek J. 1998: The Paleogene of the

Podhale Basin (Polish Inner Carpathians) – micropaleonto-
logical perspective. Przegl. Geol. 46, 721—728.

Passendorfer E. 1958: About sedimentation of the Eocene in the Tatra.

Acta Geol. Pol. 8, 451—475 (in Polish with English summary).

Perch-Nielsen K. 1985: Cenozoic calcareous nannofossils. In: Bolli

H.M., Saunders J.B. & Perch-Nielsen K. (Eds.): Plankton
stratigraphy. Cambridge University Press, Cambridge, 427—554.

Radomski A. 1967: Some stratigraphic units based on nannoplankton

in the Polish Outer Carpathians. Biul. Inst. Geol. 211, 385—393.

Radomski A. 1968: Calcareous nannoplankton zones in the Palaeo-

gene of the Western Polish Carpathians. Ann. Soc. Géol. Pol.
38, 545—605 (in Polish with English summary).

Rochon A., de Vernal A., Turon J.-L., Matthiessen J. & Head M.J.

1999: Distribution of recent dinoflagellate cysts in surface sed-
iments from the North Atlantic and adjacent seas in relation to
sea-surface parameters. Amer. Assoc. Stratigr. Palynol., Contr.
Ser. 35, 1—146.

Roniewicz P. 1969: Sedimentation of the Nummulite Eocene in the

Tatra Mts. Acta Geol. Pol. 19, 503—608 (in Polish with English
summary).

Sokołowski J. 1992: Geosynoptic documentation of the geother-

mal well Bańska IG-1. Geosynoptyka i Geotermia 1, 1—122
(in Polish).

Sokołowski S. 1973: Geology of Palaeogene and Mesozoic base-

ment of the Podhale Trough southern limb in the Column of
the Zakopane deep borehole. Biul. Inst. Geol. 265, 5—103 (in
Polish with English summary).

Soták J., Pereszlényi M., Marschalko R., Milička J. & Starek D.

2001: Sedimentology and hydrocarbon habitat of the subma-
rine-fan deposits of the Central Carpathian Paleogene Basin
(NE Slovakia). Mar. Petrol. Geol. 18, 87—114.

Soták J., Gedl P., Banská M. & Starek D. 2007: New stratigraphic

data from the Paleogene formations of the Central Western
Carpathians at the Orava region: results of integrated micropa-
leontological study in the Pucov section. Miner. Slovaca 39,
89—106 (in Slovak with English summary).

Stover L.E., Brinkhuis H., Damassa S.P., de Verteuil L., Helby R.J.,

Monteil E., Partridge A.D., Powell A.J., Riding J.B., Smelror
M. & Williams G.L. 1996: Mesozoic-Tertiary dinoflagellates,
acritarchs and prasinophytes. In: Jansonius J. & McGregor
D.C. (Eds.): Palynology: principles and applications. Amer.
Assoc. Stratigr. Palynol. Found. 2, 641—750.

Van Couvering J.A., Aubry M.-P., Berggren W.A., Bujak J.P., Naes-

er C.W. & Wieser T. 1981: The terminal Eocene event and the
Polish connection. Palaeogeogr. Palaeoclimatol. Palaeoecol.
36, 321—362.

Watycha L. 1959: Some remarks on the geology of the Podhale Fly-

sch in the eastern part of Podhale. Przegl. Geol. 8, 350—356 (in
Polish).

Wei W. & Wise S.W. Jr. 1990: Biogeographic gradients of Middle

Eocene-Oligocene calcareous nannoplankton in the South At-
lantic Ocean. Palaeogeogr. Palaeoclimatol. Palaeoecol. 79,
29—61.

Wieczorek J. 1989: The Hecho model for the Podhale Flysch?

Przegl. Geol. 9, 419—423 (in Polish).

Williams G.L., Stover L.E. & Kidson E.J. 1993: Morphology and

stratigraphic ranges of selected Mesozoic-Cenozoic dinoflagel-
late taxa in the Northern Hemisphere. Geol. Surv. Canada, Pa-
pers 92—10, 1—137.

Williams G.L., Lentin J.K. & Fensome R.A. 1998: The Lentin and

Williams index of fossil dinoflagellates, 1998 edition. Amer.
Assoc. Stratigr. Palynol., Contr. Ser. 28, 1—856.

332

GEDL and GARECKA

Appendix

An alphabetical listing of dinoflagellate cyst (their taxonomic cita-
tions are given in Williams et al. 1998) and calcareous nannoplankton
taxa found in the studied material.

Dinoflagellate cysts

Achilleodinium biformoides (Eisenack, 1954) Eaton, 1976
Achomosphaera alcicornu (Eisenack, 1954) Davey & Williams, 1966
Achomosphaera ramulifera (Deflandre, 1937) Evitt, 1963
Aiora sp. A sensu Gedl (2004a)
Amphorosphaeridium? multispinosum (Davey & Williams, 1966)

Sarjeant, 1981

Areosphaeridium diktyoplokum (Klumpp, 1953) Eaton, 1971
Areosphaeridium michoudii Bujak, 1994
Batiacasphaera micropapillata Stover, 1977
Caligodinium sp.
Charlesdowniea sp.
Cordosphaeridium gracile (Eisenack, 1954) Davey & Williams, 1966
Cordosphaeridium inodes (Klumpp, 1953) Eisenack, 1963
Cordosphaeridium minimum (Morgenroth, 1966) Benedek, 1972
Cordosphaeridium? solidospinosum Gedl, 1995
Corrudinium incompositum (Drugg, 1970) Stover & Evitt, 1978
Corrudinium? sp. A sensu Gedl (2004a)
Dapsilidinium pseudocolligerum (Stover, 1977) Bujak, Downie,

Eaton & Williams, 1980

Deflandrea phosphoritica Eisenack, 1938
Deflandrea sp.
Diphyes colligerum (Deflandre & Cookson, 1955) Cookson, 1965
Distatodinium ellipticum (Cookson, 1965) Eaton, 1976
Dracodinium laszczynskii Gedl, 1995
Dracodinium sp. A sensu Gedl (2004a)
Enneadocysta multicornuta (Eaton, 1971) Stover & Williams, 1995
Enneadocysta pectiniformis (Gerlach, 1961) Stover & Williams,

1995

Enneadocysta aff. pectiniformis sensu Gedl (2004a)
Fibrocysta bipolaris (Cookson

Eisenack, 1965) Stover & Evitt,

1978

Gongylodinium? sp. A sensu Gedl (2004)
Heteraulacacysta? leptalea Eaton, 1976
Heterosphaeridium sp. A sensu Gedl (2004a)
Homotryblium plectilum Drugg & Loeblich Jr., 1967
Homotryblium tenuispinosum Davey & Williams, 1966
Hystrichokolpoma cinctum Klumpp, 1953
Hystrichokolpoma rigaudiae Deflandre & Cookson, 1955
Impagidinium brevisulcatum Michoux, 1985
Impagidinium dispertitum (Cookson & Eisenack, 1965) Stover &

Evitt, 1978

Impagidinium sp. A sensu Gedl (2004a)
Impagidinium sp.
Lingulodinium machaerophorum (Deflandre & Cookson, 1955) Wall,

1967

Lingulodinium pycnospinosum (Benedek, 1972) Stover & Evitt, 1978
Melitasphaeridium pseudorecurvatum (Morgenroth, 1966) Bujak,

Downie, Eaton & Williams, 1980

Nematosphaeropsis lemniscata Bujak, 1984
Operculodinium centrocarpum (Deflandre & Cookson, 1955) Wall,

1967

Operculodinium aff. centrocarpum sensu Gedl (2004a)
Operculodinium? hirsutum (Ehrenberg, 1838) Lentin & Williams,

1973

Operculodinium microtriainum (Klumpp, 1953) Islam, 1983

Operculodinium tiara (Klumpp, 1953) Stover & Evitt, 1978
Pentadinium laticinctum subsp. granulatum Gocht, 1969
Rhombodinium aff. perforatum sensu Gedl (2004a)
Rhombodinium sp.
Samlandia chlamydophora Eisenack, 1954
Spiniferites pseudofurcatus (Klumpp, 1953) Sarjeant, 1970
Spiniferites ramosus (Ehrenberg, 1838) Mantell, 1854
Systematophora placacantha (Deflandre & Cookson, 1955) Davey,

Downie, Sarjeant & Williams, 1969

Thalassiphora patula (Williams & Downie, 1966) Stover & Evitt,

1978

Thalassiphora pelagica (Eisenack, 1954) Eisenack & Gocht, 1960
Tityrosphaeridium cantharellus (Brosius, 1963) Sarjeant, 1981
Wetzeliella articulata Eisenack, 1938
Wetzeliella symmetrica Weiler, 1956
Wetzeliella sp.

Calcareous nannoplankton

Braarudosphaera bigelowii (Gran & Braarud, 1935) Deflandre, 1947
Chiasmolithus expansus (Bramlette & Sullivan, 1961) Gartner, 1970
Coccolithus eopelagicus (Bramlette & Riedel, 1954) Bramlette &

Sullivan, 1961

Coccolithus formosus (Kamptner, 1963) Wise, 1973
Coccolithus pelagicus (Wallich, 1877) Schiller, 1930
Corannulus germanicus Stradner, 1962
Coronocyclus nitescens (Kamptner, 1963) Bramlette & Wilcoxon,

1967

Cribrocentrum reticulatum (Gartner & Smith, 1967) Perch-Nielsen,

1971

Cyclicargolithus floridanus (Roth & Hay, 1967) Bukry, 1971
Dictyococcites bisectus (Hay, Mohler & Wade, 1966) Bukry & Per-

cival, 1971

Dictyococcites callidus Perch-Nielsen, 1971
Dictyococcites scrippsae Bukry & Percival, 1971
Discoaster barbadiensis Tan, 1927
Discoaster binodosus Martini, 1958
Discoaster diastypus Bramlette & Sullivan, 1961
Discoaster multiradiatus Bramlette & Riedel, 1954
Discoaster saipanensis Bramlette & Riedel, 1954
Discoaster strictus Stradner, 1961
Discoaster tanii Bramlette & Riedel, 1954
Helicosphaera compacta Bramlette & Wilcoxon, 1967
Isthmolithus recurvus Deflandre, 1954
Lanternithus minutus Stradner, 1962
Neococcolithes minutus (Perch-Nielsen, 1967) Perch-Nielsen, 1971
Orthozygus aureus (Stradner, 1962) Bramlette & Wilcoxon, 1967
Pontosphaera scissura (Perch-Nielsen, 1971) Romein, 1979
Reticulofenestra dictyoda (Deflandre, 1954) Stradner, 1968
Reticulofenestra hillae Bukry & Percival, 1971
Reticulofenestra umbilica (Levin, 1965) Martini & Ritzkowski, 1968
Sphenolithus editus Perch-Nielsen, 1978
Sphenolithus moriformis (Brönnimann & Stradner, 1960) Bramlette

& Wilcoxon, 1967

Sphenolithus pacificus Martini, 1965
Sphenolithus predistentus Bramlette & Wilcoxon, 1967
Sphenolithus radians Deflandre, 1952
Sphenolithus spiniger Bukry, 1971
Thoracosphaera operculata Bramlette & Martini, 1964
Thoracosphaera saxea Stradner, 1961
Transversopontis exilis (Bramlette & Sullivan, 1961) Perch-Nielsen,

1971

Zygrhablithus bijugatus (Deflandre, 1954) Deflandre, 1959