GeolCarp_Vol52_No1_23

GEOLOGICA CARPATHICA, 52, 1, BRATISLAVA, FEBRUARY 2001

23 — 40

LATE CAMPANIAN/LATE MAASTRICHTIAN PENETRATION

OF HIGH-LATITUDE CALCAREOUS NANNOFLORA

TO THE OUTER WESTERN CARPATHIAN DEPOSITIONAL AREA

LILIAN ŠVÁBENICKÁ

Czech Geological Survey, Klárov 131/3, P.O. Box 85, 118 21 Praha 1, Czech Republic; svab@cgu.cz

(Manuscript received July 4, 2000; accepted in revised form December 12, 2000)

Abstract:  A  joint  occurrence  of  high-latitude  “boreal”  and  low-latitude  “tethyan”  calcareous  nannofossil  taxa  has
been observed in sediments of Late Campanian and Late Maastrichtian age, in the Outer (Menilite-Krosno) group of
nappes (Waschberg Zone and Ždánice Unit) and Magura group of nappes (Rača and Bílé Karpaty units) of the Outer
Western  Carpathians.  The  degree  of  influence  of  the  “boreal”  province  was  a  function  of  the  paleogeographical
position  of  basins  and  geological  time:  the  depositional  area  of  the  Outer  (Menilite-Krosno)  group  of  nappes  was
probably situated on the SE passive margin of the West European Platform. The calcareous nannofossils include both
high-latitude  species  (Monomarginatus  quaternarius,  Neocrepidolithus  watkinsii)  and  low-latitude  ones  (genera
Ceratolithoides and Quadrum) in Late Campanian times. In Maastrichtian times the high-latitude phenomena domi-
nates (common Psyktosphaera firthii, absence of Ceratolithoides kamptneri, Watznaueria : Micula ratio = 1 : 3). In the
Magura  group  of  nappes  the  depositional  area  of  which  is  considered  to  be  a  high  activity  terrane  of  the  Tethyan
mobile Realm, the Late Campanian nannoflora is characterized by the presence of low-latitude species. The influx of
high-latitude species is evident here especially in the Late Maastrichtian (higher numbers of Prediscosphaera stoveri
and  Nephrolithus  frequens).  A  migration  path  can  be  supposed  between  the  northern  part  of  the  Tethys  and  North
European  basins  through  the  Peri-Tethyan  basins  (Polish  Trough).  In  contrast,  the  Late  Campanian  section  of  the
Münsterland  Basin,  which  is  included  into  the  “boreal”  province,  yielded  exclusively  cosmopolitan  and  boreal
nannofossils including Heteromarginatus bugensis.

Key words: Outer Western Carpathians, Münsterland Basin, Campanian, Maastrichtian, calcareous nannofossils,
biostratigraphy, bioprovinces.

Introduction

Bioprovincialism was increasing during the Late Cretaceous
due to gradual global cooling, resulting in the coexistence of
typical  Tethyan/tropical  and  Boreal  or  Austral  fauna.  Ac-
cording  to  Burnett  (1998),  paleobiogeographical  separation
of  calcareous  nannofloras  became  increasingly  apparent
through the Campanian and into the Maastrichtian. This pa-
leobiogeographic separation allows for the differentiation of
three distinctive biogeographical regions: the “boreal” prov-
ince  (broadly,  northern  high  paleolatitudes),  the  intermedi-
ate- “tethyan” province (broadly, moderate to low paleolati-
tudes),  and  the  “austral”  province  (southern  high
paleolatitudes).

A joint occurrence of low-latitude “tethyan” and high-lati-

tude  “boreal”  calcareous  nannofossil  taxa  was  observed  in
Late Cretaceous sediments of the Outer Western Carpathians
(Figs.  1  and  2),  particularly  in  the  Outer  (Menilite-Krosno)
group of nappes (Waschberg Zone and Ždánice Unit) and in
the Magura group of nappes (Rača and Bílé Karpaty units).
The depositional areas of these geological units are supposed
to have originated south-east of their present locations and,
as  a  whole,  they  may  be  included  in  the  warm-temperate
realm  of  the  Tethys.  According  to  Stráník  et  al.  (1996)  the
depositional  area  of  the  Outer  (Menilite-Krosno)  group  of
nappes was situated about 100 km from its present location,
on the southeastern passive margin of the European Platform
which  is  believed  to  be  a  northern  margin  of  the  Tethys.

Stráník (in Švábenická et al. 1997) regards the Magura group
of nappes depositional area as a high activity terrane of the
Tethyan mobile Realm, situated a few hundreds km from its
present location, partially on the territory of the present Cen-
tral Carpathians and touching the southeastern rim of Euro-
pean Platform.

Fig. 1. A sketch map showing the locations of studied areas (marked
by hatched areas).

24 ŠVÁBENICKÁ

Autochthonous epicontinental Cretaceous sediments of the

cool to temperate “boreal” realm, that is the chalk shelf depo-
sitional area of North European basins, occur farther to the
north and northwest in the European Platform.

The aim of this work was to study the migration of high-lati-

tude calcareous nannoflora to the depositional area of the
northern margin of Tethys during the Campanian and Maas-
trichtian, and influence of the “boreal” province in the particu-
lar depositional areas in the framework of the present Outer
Western Carpathians.

Previous studies

The presence of distinctive latitude-restricted nannoplank-

ton species was first reported by Worsley & Martini (1970)
in  the  Maastrichtian,  who  noted  the  bipolar  distribution  of
Nephrolithus frequens and a coeval concentration of Micula
murus  in  lower  latitudes.  Bukry  (1973)  proposed  low-lati-
tude nannoplankton biostratigraphic zonation for the Upper
Cretaceous. Latitude and paleoclimatic distribution patterns
of  Maastrichtian  nannoflora  were  discussed  by  Worsley
(1974).  Thierstein  (1976)  noted  that  some  nannofossil  spe-
cies  display  high  abundances  with  increasing  paleolatitude,
and Thierstein & Haq (1977) presented a systematic analysis
of the Upper Cretaceous nannoplankton biogeography. Many
later  publications  discussed  latitude-restricted  nannofossils
and their affinities to provinces during the Upper Cretaceous:
Wind  (1979),  Perch-Nielsen  (1985),  Burnett  (1990,  1998),
Watkins (1992), Watkins et al. (1996), and others.

The coexistence of low-latitude and high-latitude nanno-

fossils in the Western Carpathians was first observed by

Švábenická (1992) in Campanian sediments of the Pavlov-5
borehole  (see  Fig.  3)  in  the  Ždánice  Unit,  Outer  (Menilite-
Krosno) group of nappes. Later, this phenomenon was also
observed in the Campanian and Maastrichtian flysch depos-
its of the Magura group of nappes in the Rača and Bílé Kar-
paty  units  (Švábenická  et  al.  1997;  Švábenická  in  Bubík  et
al. 1999). The significant penetration of high-latitude nanno-
flora to the depositional area of the Outer (Menilite-Krosno)
group of nappes (Waschberg Zone and Ždánice Unit) in the
Turonian—Maastrichtian  interval  was  broadly  described  by
Švábenická  (in  Stráník  et  al.  1996;  in  Summesberger  et  al.
1999; in Hofmann et al. 1999). A stratigraphic correlation of
the  latitude-restricted  nannofossils  in  the  Campanian  of  the
Outer  Western  Carpathians  was  presented  by  Švábenická
(1995 and 1998). High-latitude “boreal” species were found
to be more frequent in the Outer group of nappes than in the
Magura group of nappes (Švábenická 1999). An overview of
the occurrences of high- and low-latitude species in nanno-
fossil  assemblages  within  the  depositional  area  of  the  Late
Cretaceous northern margin of the Tethys was presented by
Švábenická et al. (in print).

Material

Material was collected in the Outer Western Carpathians in

the  territory  of  the  Czech  Republic,  Slovakia  and  Austria
(Figs.  3  and  4).  Data  presented  here  were  obtained  mostly
during  geological  research  in  this  area  and  are  mostly  pub-
lished. This work summarizes all available observations.

Samples were taken from outcrops: from flysch sediments

s.s. (interval T

sensu Bouma 1962) in the Rača and Bílé

Fig. 2. Schematic geological map of the Alps, Carpathians, Dinarides and the Pannonian Basin, and the location of the study area. From
Linzer (in Tomek 1996), modified.

CAMPANIAN/MAASTRICHTIAN PENETRATION OF HIGH-LATITUDE NANNOFLORA 25

Karpaty units, Magura group of nappes, and from claystones
and marlstones in the Waschberg Zone and Ždánice Unit,
Outer (Menilite-Krosno) group of nappes. In addition, tecton-
ic slices of the Púchov Marl in the front of the Magura group
of nappes were studied. These sediments are believed to have
originated in the depositional area of the Pieniny Klippen
Belt (Švábenická et al. 1997).

Outer Western Carpathians

Outer (Menilite-Krosno) group of nappes

Waschberg Zone and Ždánice Unit

Pálava Formation (Stráník et al. 1996): grey and brownish-

grey calcareous claystones with a variable amount of silt.
Nannofossils were studied in the Pavlov-5 borehole (Švábe-
nická 1992) and at the localities of Mikulov and Děvín
(Stráník et al. 1996), Ernstbrunn (Hofmann et al. 1999), East

Michelstetten and West Klement (Summesberger et al.
1999). In total, 60 samples were studied.

Magura group of nappes (western part)

Rača Unit

Kaumberg Formation: the upper part is formed by medium-

to thin-bedded flysch with sporadic layers of red-brown cal-
careous claystones to marlstones. Nannofossils were studied
at the locality of Salajka (Švábenická et al. 1997). Four sam-
ples were studied.

Soláň Formation (Matějka & Roth 1948): medium- to

thick-bedded flysch. Its Cretaceous sediments are represent-
ed by blue-grey, fine- to coarse-grained sandstones prevail-
ing over grey and dark grey-green claystones. Nannofossils
were studied at the localities of Valašské Meziříčí (Šváben-
ická et al. 1997 and see Appendix 1.4), Racková (see Appen-
dix 1.3) and Uzgruň (Bubík et al. 1999). In total, 22 samples
were studied.

Fig. 3. A sketch map of the western part of the Outer Western Carpathians showing the locations of studied sections (in capitals) and ma-
jor cities. Pv-5: Pavlov-5 borehole.

26 ŠVÁBENICKÁ

Bílé Karpaty Unit

Ondrášovec Member (Potfaj 1993) of the Kaumberg For-

mation: thin-bedded flysch with red-brown calcareous clay-
stones. This lithofacies is developed in the upper part of the
Kaumberg Formation. Nannofossils were studied at the locali-
ties of Janegov Mlyn and Velký Lopeník. In total, 13 samples
were studied.

Javorina Formation (Stráník et al. 1989): fine- to medium-

bedded  flysch  contains  fine-  to  medium-grained  greywacke
sandstones,  mostly  non-calcareous  grey  claystones  and  spo-
radically  intercalations  of  turbidity  limestones.  Nannofossils
were  studied  at  the  localities  of  Klokočník  (Švábenická  &
Bubík 1992) and Javořina (see Apendix 1.2). In total, 30 sam-
ples were studied.

Svodnice Formation (Pesl 1968): medium- to thick-bedded

flysch with grey calcareous claystones prevailing over
greywacke sandstones. Nannofossils were studied at the local-

ity of Štítná (Švábenická et al. 1997). Twenty samples were
studied.

A tectonic slice (Púchov Marl) in the front of the Bílé Kar-

paty Unit is characterized by red, highly calcareous claystones
and marls. Sediments were studied at the locality of Hluk (see
Appendix 1.1). Five samples were studied.

Münsterland Basin, NW Germany

Upper Campanian nannofossil associations from the bore-

holes  of  Oberdarfeld-1  (Švábenická  in  Kaever  &  Lom-
merzheim  1995)  and  Longinusturm-1  (Lommerzheim  1995)
were included in this study to provide a comparison with co-
eval associations from sediments deposited on the West Euro-
pean  Platform.  This  depositional  area  is  included  within  the
“boreal” province (Figs. 1 and 5). In total, 150 samples were
studied.

Fig. 4. Chronostratigraphic correlation chart of the lithostratigraphic units of the Outer Western Carpathians mentioned in the text.
* Svodnice Formation: medium- to thick-bedded flysch with grey calcareous claystones.

OUTER WESTERN CARPATHIANS

OUTER GROUP OF

NAPPES

MAGURA GROUP OF NAPPES

sil z

one

sing

h (

197

ch-

Nie

lse

n (

85)

sil z

one

tt (

98)

WASCHBERG ZONE

ŽDÁNICE UNIT

RAČA UNIT

BÍLÉ

KARPATY

UNIT

TECTONIC

SLICES of

uncertain origin

CC26

UC2

UC1

MAAS

UC1

CC22
CC21
CC20

UC1

CC19

ANIA

CC18

UC1

Pálava Formation

grey and brownish-grey

calcareous claystones with a

variable amount of silt

Soláň

Formation

medium- to

thick-bedded

flysch with grey

and dark grey-

green

claystones

Kaumberg

Formation

medium- to

thin-bedded

flysch with rare

red-brown
calcareous
claystones

Svodnice
Form.

Javorina

Formation

fine- to

medium-bedded

flysch

with grey

claystones

Ondrá-
šovec
Member

thin-bedded

flysch with red-

brown calc.

claystones

Kaumberg

Formation

Púchov Marl

red, highly

calcareous

claystones and

marls

28 ŠVÁBENICKÁ

and Ždánice Unit. The specimens were slightly etched and
broken only rarely, and therefore easily identifiable.

The abundance, species diversity and preservation of nanno-

fossil assemblages in the Longinusturm-1 borehole depend on
the lithological character of the sediment. Specimens from
sandstones and limestone intercalations were poorly pre-
served, broken, etched or overgrown with calcite; identifica-
tion of some specimens was therefore difficult. Marlstones and

claystones provided moderately well preserved, easily identifi-
able specimens.

Methods

Suspension slides were prepared using a decantation meth-

od. The 3—30 µm fraction was separated in the following way:

Fig. 7. Calcareous nannofossil distribution at the localities of Janegov Mlyn and Velký Lopeník, Outer Western Carpathians (Bílé Kar-
paty Unit, Ondrášovec Member of the Kaumberg Formation). Estimated abundance of nannofossil taxa: A = abundant (>5 specimens per
field of view), C = common (5—1 specimens per field of view),  F  =  few  (>1  specimen  per  10  field  of  view),  R  =  (<1  specimen  per  10
fields  of  view),  r  =  reworked  species,  f  =  fragments.  Preservation  of  nannofossils:  M  =  moderate  (overgrowth,  etching  or  mechanical
damage  is  apparent  but  majority  of  specimens  are  easily  identifiable),  P  =  poor  (etching  and  mechanical  damage  is  intensive  making
identification of some specimes difficult), VP = very poor (some specimens cannot be  identified). Estimates of the abundance of nanno-
fossils in samples: H = high (>5 nannofossils per field of view), M = moderate (1—5 nannofossils per field of view), L = low (<1 nanno-
fossil per field of view). Categories after Burnett & Whitham (1999), modified.

CAMPANIAN/MAASTRICHTIAN PENETRATION OF HIGH-LATITUDE NANNOFLORA 29

the heavy-fraction was allowed to settle for 3 minutes in a
45 mm water column and removed, the fine-fraction was
saved for slide preparation after 45 minutes. The slides were
studied using a Nikon light-microscope at 1000

magnifica-

tion.

Quantitative and other data of the nannofossil taxa men-

tioned in this study were collected using the method of Burnett
& Whitham (1999) and slightly modified. The calculation of

relative abundances of nannofossil taxa was based on ca. 300—
500 specimens. Biostratigraphic data were correlated with the
UC zones introduced by Burnett (1998) and compared with
the standard nannoplankton CC zones by Sissingh (1977) and
Perch-Nielsen (1985). Interpretations of provincial preferences
of nannofossil species were based on Wind (1979), Watkins et
al. (1996) and Burnett (1998). The herein used terms “boreal”
and “tethyan” follow Burnett (1998).

Fig. 8. Calcareous nannofossil distribution at the locality of Štítná, Outer Western Carpathians (Bílé Karpaty Unit, Svodnice Formation).
Estimated abundance of nannofossil taxa: C = common (5—1 specimens per field of view), F = few (>1 specimen per 10 fields of view), R
= (<1 specimen per 10 fields of view), ? = questionable species. Preservation of nannofossils: M = moderate (overgrowth, etching or me-
chanical damage is apparent but majority of specimens are easily identifiable), P = poor (etching and mechanical damage is intensive
making identification of some specimens difficult), VP = very poor (some specimens cannot be identified). Estimates of the abundance of
nannofossils in samples: H = high (>5 nannofossils per field of view), M = moderate (1—5 nannofossils per field of view), L = low (<1
nannofossil per field of view). Categories after Burnett & Whitham (1999), modified.

32 ŠVÁBENICKÁ

Results

Outer Western Carpathians

Outer (Menilite-Krosno) group of nappes, Waschberg Zone
and Ždánice Unit

Campanian, zones UC15c

—UC15d

; UC15d

(CC21—

CC22)

Nannofossil associations are characterized by high species

diversity and by the presence of both low- and high-latitude
species. Low-latitude nannofossils are represented by genera
Uniplanarius and Ceratolithoides, high-latitude species by
genera Biscutum (small and large species), Monomarginatus,
and species Neocrepidolithus watkinsii and Acuturris scotus.
Specimens of genus Lucianorhabdus and species Kamptnerius
magnificus represent up to 5 % of the assemblages. Watznaue-
ria barnesae quantitatively prevails over Micula staurophora
in the approximate ratio 2 : 1.

Upper Maastrichtian, zone UC20d

(CC26)

The highly diversified assemblage is characterized by high

numbers  of  species  Arkhangelskiella  cymbiformis,  Micula
staurophora  (Watznaueria : Micula  ratio  =  1 : 3),  Predis-
cosphaera ex gr. cretacea, and Cribrosphaerella ehrenbergii
forming 5—10 % of the assemblages, and by common (1—5 %)
Lithraphidites quadratus, Prediscosphaera majungae, and Ah-
muellerella regularis. Low-latitude “tethyan” species are rep-
resented  by  rare  Micula  murus  and  M.  prinsii,  high-latitude
species  by  Nephrolithus  frequens,  Prediscosphaera  stoveri
and  common  Psyktosphaera  firthii.  Moreover,  Biantholithus
sparsus was recorded here on rare occasion. Specimens of ge-
nus Ceratolithoides were not observed.

Magura group of nappes, Rača and Bílé Karpaty units

Campanian and basal Maastrichtian, zones UC15b

—UC16

(CC20—CC23a)

Nannofossil assemblages are rather of “tethyan” affinity,

with only minor amounts of high-latitude species (Figs. 6 and
7).  The  associations  are  characterized  by  higher  numbers  of
Watznaueria barnesae, Cribrosphaerella ehrenbergii and Pre-
discosphaera cretacea. A usual component of the assemblages
are  low-latitude  taxa  including  Ceratolithoides  (C.  aculeus
prevails), and Uniplanarius sissinghii and U. trifidus. Though
infrequent,  high-latitude  species,  such  as  Biscutum  coronum
and  Prediscosphaera  stoveri,  are  also  present.  Neocrepido-
lithus watkinsii was recorded in the Rača Unit (Valašské Mezi-
říčí  locality  –  see  Appendix  1),  in  the  assemblage  where
Watznaueria : Micula ratio was observed of approx. 1 : 1.

A tectonic slice in the front of the Bílé Karpaty Unit

(Púchov Marl) was studied at Hluk locality (see Appendix 1).
The  nannofossil  assemblage  consists  of  high  numbers  (5—
10 %) of Watznaueria barnesae and Cribrosphaerella ehren-
bergii;  Lucianorhabdus  is  absent.  Uniplanarius  trifidus,  U.
sissinghii,  Ceratolithoides  aculeus,  Micula  staurophora  and

Reinhardtites levis are relatively common, representing ca. 1—
3 % of the assemblages. Biscutum coronum, B. ellipticum,
Prediscosphaera stoveri, Kamptnerius magnificus, Petrarhab-
dus copulatus and Tortolithus cf. hallii were recorded in very
low numbers (< 0.1 %).

Upper Maastrichtian, zone UC20 (CC26)

Nannofossil assemblages of relatively high species diver-

sity  (Fig.  8)  include  Lithraphidites  quadratus,  Eiffellithus
paralleus, Markalius apertus and Corollithion ?madagaska-
rensis. The assemblages are generally characterized by high-
er  numbers  of  the  species  Arkhangelskiella  cymbiformis,
Watznaueria  barnesae  and  Micula  staurophora  (forming
about 5 %), and by the influx of high-latitude species. This is
confirmed  by  the  presence  of  Prediscosphaera  stoveri,  Bis-
cutum  coronum,  fragments  of  Cribrosphaerella  cf.  daniae,
rare Acuturris scotus and very rare Psyktosphaera firthii. Low-
latitude species are represented by Ceratolithoides kamptneri,
Micula murus and M. prinsii. The Watznaueria : Micula ratio
is  ca.  1 : 2.  Biantholithus  sparsus  was  also  recorded  here  on
rare occasion.

Münsterland Basin

Sediments of the Longinusturm-1 borehole provided nan-

nofossil assemblages of cosmopolitan and high-latitudinal
affinities (Fig. 9) indicating a Late Campanian age (zone
UC15d-e

). An exception is sample 59.50 m where one

specimen of  Uniplanarius trifidus (see Fig. 12.36) and rare
Cylindralithus serratus (see Fig. 12.29,30) were found. As-
semblages  are  characterized  by  a  high  number  of  Lucia-
norhabdus,  and  a  low  number  of  Cribrosphaerella  ehren-
bergii.  This  phenomenon  was  especially  evident  in  the
interval of 31—38 m.

The whole section contains species Eiffellithus eximius,

Broinsonia parca constricta and relative common Reinhard-
tites  anthophorus  and  R.  levis.  Small  specimens  of  Predis-
cosphaera stoveri were recorded already in the lowermost part
of  the  section,  in  association  with  Orastrum  campanensis,
Heteromarginatus  bugensis,  Neocrepidolithus  watkinsii,  Bis-
cutum coronum, B. dissimilis, Acuturris scotus and rare Qua-
drum svabenickae. Rare occurrences of Petrarhabdus copula-
tus  were  observed  only  occasionally,  in  the  interval  of
37.80—88.40 m. Nannofossil associations of the Upper Cam-
panian  sediments  in  the  borehole  Oberdarfeld-1  provided
similar  data;  no  “tethyan”  nannofossils  were  recorded  here
(Švábenická in Kaever & Lommerzheim 1995).

The occurrences of latitude-restricted nannofossil species

in the above mentioned geological units are presented in
Figs. 10 and 11.

Discussion

According to Kaever & Lommerzheim (1995), the Münster-

land Basin had a transitional position between the Boreal and
Tethyan realms in the Late Campanian. This is supported by

CAMPANIAN/MAASTRICHTIAN PENETRATION OF HIGH-LATITUDE NANNOFLORA 33

Fig. 10. Upper Campanian. Occurrence of latitude-restricted nannofossils in the Outer Western Carpathians and the Münsterland Basin.
C = common (>1 specimens per field of view), F = few (>1 specimen per 10 fields of view), R = rare (<1 specimen per 10 fields of view),
VR = specimen recorded on very rare occasions, - = specimen not recorded.

the  occurrence  of  rare  “tethyan”  and  cosmopolitan  taxa  and
demonstrated by the Late Campanian sequence of the Ober-
darfeld section. It has important implications for the strati-
graphical  correlation  between  the  Boreal  Realm  and  the
Tethys: cephalopods Nostoceras polyplocum, Trochoceramus
aff.  helveticus  and  Belemnitella  mucronata  minor  coexist
here,  nevertheless,  nannofloras  include  “boreal”  index  spe-
cies only. Similar nannofossil associations were observed in
the Longinusturm-1 borehole. The absence of typical “tethy-
an” nannofossils, such as Ceratolithoides and Uniplanarius,
within  zone  UC15d-e

supports the idea that nannofloras

from the Tethys did not penetrate into this depositional area.
The  only  minor  influx  of  low-latitude  nannoflora  into  the
NW part of the Münsterland Basin was recorded at a depth of
59.50  m  where  rare  Cylindralithus  serratus  and  a  single
specimen  of  Uniplanarius  trifidus  were  observed.  The  ab-
sence of low-latitude nannofossil species does not allow us
to use the nannoplankton biozonation for the “tethyan-inter-
mediate province” sensu Burnett (1998) for the Upper Cam-
panian sediments in the Münsterland Basin.

Burnett (1998) presented Heteromarginatus bugensis and

Neocrepidolithus watkinsii as apparently endemic taxa re-

stricted to high paleolatitudes, and Kamptnerius, Nephrolithus
and large Biscutums as taxa which, although globally distribut-
ed, become more abundant in high latitudes. The presence and
higher numbers of the above mentioned nannofossils are clear
evidence  of  some  influence  of  the  “boreal”  province  in  the
Waschberg Zone and Ždánice Unit during the Campanian and
Maastrichtian. According to Stráník et al. (1996) this deposi-
tional area was situated on the SE passive margin of West Eu-
ropean  Platform  and  belonged  to  the  Boreal-Tethyan  transi-
tional area. Heteromarginatus bugensis has not been observed
in  the  Outer  Western  Carpathians  and  was  recorded  in  the
Münsterland Basin only.

The gradual change in the character of nannoplankton as-

semblages is apparent especially in the Outer (Menilite-Kros-
no) group of nappes (Waschberg Zone and Ždánice Unit) from
the  Campanian  to  Maastrichtian.  The  co-occurrence  of  low-
and high-latitude nannofossils is obvious in the Late Campa-
nian  whereas  high-latitude  taxa  become  more  abundant  and
low-latitude  taxa  nearly  disappear  in  the  Late  Maastrichtian
(absence of Ceratolithoides during the Maastrichtian). These
data  may  indicate  gradual  cooling  within  this  depositional
area,  probably  caused  by  both  the  increasing  dominance  of

34 ŠVÁBENICKÁ

southerly paleocurrents from the North European basins and
the global cooling during the Late Cretaceous. In addition, the
detrital and nutrient input and other factors have to be also tak-
en into account.

The joint occurrence of latitude-restricted nannofossils in

the Outer (Menilite-Krosno) group of nappes (Waschberg
Zone and Ždánice Unit) documents that the depositional area
of the northern margin of the Tethys was influenced by the

Fig.  12.  Stratigraphically  significant  calcareous  nannofossils  from  the  Outer  Western  Carpathians  and  Münsterland  Basin.  PPL:  plane-
polarized  light,  XPL:  cross-polarized  light.    1,  2.    Heteromarginatus  bugensis  (Górka)  Crux;  Upper  Campanian,  Münsterland  Basin,
Longinusturm-1, 88.40 m; 1, PPL; 2, XPL.  3, 4.  Orastrum campanensis (Čepek) Wind & Wise; Upper Campanian, Outer Western Car-
pathians, Waschberg Zone, Ernstbrunn  No. 6064; 3, PPL; 4, XPL.  5, 6. Biscutum magnum Wind & Wise; Upper Campanian, Outer West-
ern Carpathians, Waschberg Zone, Ernstbrunn No. 6064; 5, PPL; 6, XPL.  7, 8.  Biscutum melaniae (Górka) Burnett; Upper Campanian,
Münsterland Basin, Longinusturm-1, 37.80 m; 7, PPL; 8, XPL.  9, 10.   Biscutum ellipticum (Górka) Grün; Upper Campanian, Münster-
land Basin, Longinusturm-1, 88.40 m; 9, PPL; 10, XPL.  11, 12.  Biscutum coronum Wind & Wise; Upper Campanian, Münsterland Ba-
sin, Longinusturm-1, 37.80 m; 11, PPL; 12, XPL. 13, 14.  Neocrepidolithus cohenii (Perch-Nielsen) Perch-Nielsen; Upper Maastrichtian,
Outer Western Carpathians, Magura group of nappes, Uzgruň No. 21A; 13, PPL; 14, XPL.  15—18.  Neocrepidolithus watkinsii Pospichal
& Wise; Upper Campanian; 15,16: Münsterland Basin, Longinusturm-1, 31.70 m.   17, 18: Outer Western Carpathians, Waschberg Zone,
Ernstbrunn No. 6064; 15,17, PPL; 16, 18, XPL.  19, 20.  Reinhardtites anthophorus (Deflandre) Perch-Nielsen; Upper Campanian, Outer
Western  Carpathians,  Waschberg  Zone,  Ernstbrunn  No.  6064;  19,  PPL;  20,  XPL.   21,  22.   Reinhardtites  levis  Prins  &  Sissingh;  Upper
Campanian, Münsterland Basin, Longinusturm-1, 88.40 m; 21, PPL; 22, XPL.  23, 24.  Monomarginatus quaternarius Wind & Wise; Up-
per Campanian, Outer Western Carpathians, Waschberg Zone; 23: Děvín; 24: Ernstbrunn No. 6064; XPL.  25—27. Petrarhabdus copula-
tus (Deflandre) Wind & Wise; Upper Campanian, Outer Western Carpathians; 25: tectonic slice, Púchov Marl, Hluk; 26, 27: Wachberg
Zone,  Ernstbrunn  No.  6064;  25,  26,  PPL;  27,  XPL.    28—30.  Cylindralithus  serratus  Bramlette  &  Martini;  Upper  Campanian;  28:  side
view, Outer Western Carpathians, Púchov Marl (tectonic slice in the Magura group of nappes), Hluk; 29, 30: Münsterland Basin, Longi-
nusturm-1, 31.70 m; 28, 29, PPL; 30, XPL.  31, 32.  Angulofenestrellithus  snyderi Bukry; Upper Campanian, Outer Western Carpathians,
Waschberg  Zone,  Ernstbrunn  No.  6064;  31,  PPL;  32,  XPL.    33,  34.    Uniplanarius  sissinghii  Perch-Nielsen;  Upper  Campanian,  Outer
Western  Carpathians,  Waschberg  Zone,  Ernstbrunn  No.  6064;  33,  PPL;  34,  XPL.    35,  36.    Uniplanarius  trifidus  (Stradner)  Hattner  &
Wise;  Upper  Campanian;  35:  Outer  Western  Carpathians,  Waschberg  Zone,  Ernstbrunn  No.  6064;  36:  Münster  Basin,  Longinusturm-1,
59.50  m;  35,  PPL;  36,  XPL.    37,  38.    Podorhabdus?  elkefensis  Perch-Nielsen;  Upper  Maastrichtian,  Outer  Western  Carpathians,  Štítná
No.  35C;  37,  PPL;  38,  XPL.    39—42.  Arkhangelskiella  cymbiformis  Vekshina;  Outer  Western  Carpathians;  39—40:  Upper  Campanian,
Waschberg  Zone,  Ernstbrunn  No.  6064;  41—42:  Upper  Maastrichtian,  Magura  group  of  nappes,  Uzgruň  No.  26A

; 39, 41, PPL; 40, 42,

XPL. Microphotographs by L. Švábenická.

▲

Fig. 11. Upper Maastrichtian. Occurrence of latitude-restricted nannofossils in the Outer Western Carpathians. For quantitative explana-
tions see Fig. 10.

CAMPANIAN/MAASTRICHTIAN PENETRATION OF HIGH-LATITUDE NANNOFLORA 37

Fig. 13: Stratigraphically significant calcareous nannofossils from the Outer Western Carpathians and Münsterland Basin. PPL: plane-polar-
ized light, XPL: cross-polarized light. 1, 2. Ceratolithoides kamptneri Bramlette & Martini; Upper Maastrichtian, Outer Western Carpathians,
Magura group of nappes, Uzgruň 26A

; 1, PPL; 2, XPL. 3, 4. Ceratolithoides sesquipedalis Burnett; Upper Campanian, Outer Western Car-

pathians, Waschberg Zone, Ernstbrunn No. 6064; 3, PPL; 4, XPL. 5. Ceratolithoides aculeus (Stradner) Prins & Sissingh; Upper Campanian—
?basal  Maastrichtian,  Outer  Western  Carpathians,  Púchov  Marl  (tectonic  slice  in  the  Magura  group  of  nappes),  Hluk;  XPL.  6.  Corollithion
?madagaskarensis Perch-Nielsen; Upper Maastrichtian, Outer Western Carpathians, Magura group of nappes, Štítná No. 35C; XPL. 7—10. Cri-
brocorona gallica (Stradner) Perch-Nielsen; Upper Maastrichtian, Outer Western Carpathians; 7, 8: Magura group of nappes, Štítná No. 29F;
9, 10: Waschberg Zone, West Klement; 7, 9, PPL; 8, 10, XPL. 11, 12.  Nephrolithus frequens Górka; Upper Maastrichtian, Outer Western Car-
pathians, Magura group of nappes, Uzgruň 21; 11, PPL; 12, XPL. 13, 14. Eiffellithus eximius (Stover) Perch-Nielsen; Upper Campanian, Outer
Western Carpathians,  Waschberg  Zone, Michelstetten; 13, PPL; 14, XPL.  15,  16. Eiffellithus parallelus Perch-Nielsen; Upper Maastrichtian,
Outer Western Carpathians, Waschberg Zone, West Klement, 15, PPL; 16, XPL. 17, 18. Ahmuellerella regularis (Górka) Reinhardt & Górka;
Upper Maastrichtian, Outer Western Carpathians, Magura group of nappes, Uzgruň No. 26A

; 17, PPL; 18, XPL. 19, 20. Micula staurophora

(Gardet) Stradner; Upper Maastrichtian, Outer Western Carpathians, Magura group of nappes, Štítná No. 35C; 19, PPL; 20, XPL. 21, 22.  Micu-
la  prinsii  Perch-Nielsen;  Upper  Maastrichtian  Outer  Western  Carpathians,  Waschberg  Zone,  West  Klement;  21,  PPL;  22,  XPL.  23,  24.  Tet-
rapodorhabdus  decorus  (Deflandre)  Wind  &  Wise;  Upper  Maastrichtian,  Outer  Western  Carpathians,  Waschberg  Zone,  West  Klement;  23,
PPL; 24, XPL.  25, 26.   Prediscosphaera majungae Perch-Nielsen (proximal shield and side view); Upper Maastrichtian, Outer Western Car-
pathians, Waschberg Zone, West Klement; 25, PPL; 26, XPL. 27, 28. Prediscosphaera stoveri (Perch-Nielsen) Shafik & Stradner; Upper Maas-
trichtian, Outer Western Carpathians; 27: Waschberg Zone, West Klement; 28: Magura group of nappes, Uzgruň No. 20B; XPL. 29, 30. Predis-
cosphaera arkhangelskyi (Reinhardt) Perch-Nielsen; Upper Campanian, Outer Western Carpathians, Waschberg Zone, Michelstetten; XPL. 31.
Lithraphidites quadratus Bramlette & Martini; Upper Maastrichtian, Outer Western Carpathians, Magura group of nappes, Štítná No. 29F; PPL.
32,  33.    Cribrosphaerella  ehrenbergii  (Arkhangelsky)  Deflandre;  Münsterland  Basin,  Longinusturm-1,  37.80  m;  32,  PPL;  33,  XPL.  34—36.
Psyktosphaera firthii Pospichal & Wise; Upper Maastrichtian, Outer Western Carpathians, Waschberg Zone, West Klement; 34, PPL; 35, 36,
XPL. 37, 38. Markalius inversus (Deflandre) Bramlette & Martini; Upper Maastrichtian, Outer Western Carpathians, Magura group of nappes,
Štítná No. 29F; 37, PPL; 38, XPL. 39, 40.  Markalius apertus  Perch-Nielsen; Upper Maastrichtian, Outer Western Carpathians, Magura group
of  nappes,  Uzgruň  No.  21;  39,  PPL;  40,  XPL.  41,  42.  Biantholithus  sparsus  Bramlette  &  Martini;  Upper  Maastrichtian,  Outer  Western  Car-
pathians, Magura group of nappes, Uzgruň No. 20; 41, PPL; 42, XPL. Microphotographs by L. Švábenická.

Boreal region in the Late Cretaceous. The existence of a cor-
ridor can be supposed across the Peri-Tethyan basins (Polish
Trough)  probably  connecting  the  northern  Tethyan  basins
with  North  European  basins  from  the  Turonian—Coniacian
until  the  Maastrichtian—Paleocene  (Malata  &  Poprawa
1997). This path could have caused the migration of high-lat-
itude “boreal” nannoflora into depositional areas of the Outer
Western  Carpathians  during  the  Campanian—Maastrichtian
interval. The abundance of low-latitude taxa and the different
character of nannofossil assemblages in the Magura group of
nappes document, that its depositional area was located far to
the  south  and  was  separated  from  those  of  the  Waschberg-
Ždánice  Unit  by  the  Silesian  Basin  and  Silesian  Cordillera
(Potfaj 1998).

The presence of the Paleogene marker species Biantholithus

sparsus in nannofossil assemblages otherwise Late Maastrich-
tian  in  age  is  mentioned  from  time  to  time,  for  instance  by
Heck & Prins (1987) and Pospichal & Wise (1990). This spe-
cies  is  occasionally  observed  in  small  numbers  in  the  Maas-
trichtian  sediments  of  the  Outer  Western  Carpathians,  in  the
Pálava Formation of the Waschberg Zone, Austria (Summes-
berger  et  al.  1999)  and  the  Soláň  Formation  of  the  Magura
group of nappes (Bubík et al. 1999). The stratigraphic signifi-
cance of B. sparsus seems to be therefore arguable and should
be subject to a detailed revision.

Conclusions

The common occurrence of high- and low-latitude nanno-

fossils in the Outer (Menilite-Krosno) group of nappes
(Waschberg Zone and Ždánice Unit) documents that the dep-

ositional area of the northern margin of the Tethys was con-
nected with the North European basins in the Campanian and
Maastrichtian, probably through a corridor of Peri-Tethyan
basins, Polish Trough.

The degree of influence by “tethyan” and “boreal” bio-

provinces was a function of both the geographical positions
of the basins and time:

In the Upper Campanian, “boreal” species (Neocrepido-

lithus watkinsii, Monomarginatus quaternarius, Acuturris
scotus, Biscutum coronum) form a significant component of
the nannofossil assemblages in the Outer (Menilite-Krosno)
group of nappes (Waschberg Zone and Ždánice Unit). In con-
trast, they are observed in small numbers or are absent in the
Magura group of nappes.

In the Upper Maastrichtian, a significant input of high-lati-

tude  nannoflora  is  evident  in  the  Outer  (Menilite-Krosno)
group  of  nappes  (Waschberg  Zone  and  Ždánice  Unit).  As
concerns  the  Magura  group  of  nappes,  gradual  “boreal”  in-
fluence  was  recorded  here,  nevertheless,  the  nannoflora  re-
tains its “tethyan“ character.

In the Upper Campanian, similar cosmopolitan and high-

latitude  nannofossil  species  occur  in  the  Outer  (Menilite-
Krosno)  group  of  nappes  (Waschberg  Zone  and  Ždánice
Unit)  and  Münsterland  Basin.  An  exception  is  the  species
Heteromarginatus bugensis which was observed exclusively
in the Münsterland Basin.

Low-latitude nannofossil species are nearly absent from

the Upper Campanian sediments of the Münsterland Basin.

By virtue of the common occurrence of high- and low-lati-

tude nannofossil species, both “boreal” and “tethyan” zona-
tions sensu Burnett (1998) can be applied to biostratigraphic
correlations of the Campanian and Maastrichtian deposits in

▲

38 ŠVÁBENICKÁ

the Outer (Menilite-Krosno) group of nappes of the Outer
Western Carpathians.

Acknowledgements:  This  study  is  a  contribution  to  the
project  “Low-  and  high-latitude  nannofossils  and  their  bios-
tratigraphic  significance  (Campanian  and  Maastrichtian)“,
supported by the Grant Agency of the Czech Republic (Grant
No. 205/97/0687). The author thanks Prof. Dr. Matthias Kaev-
er (Westfälische Wilhelms Universität Münster) for providing
the study material from the Münsterland Basin and Dr. Zdeněk
Stráník (Czech Geological Survey) for the fruitful discussion.

References

Bouma A.H. 1962: Sedimentology of some flysch deposits; a

graphic approach to facies interpretation. Elsevier, 1—168.

Bubík M., Bąk M. & Švábenická L. 1999: Biostratigraphy of the

Maastrichtian to Paleocene distal flysch sediments of the
Rača Unit in the Uzgruň Section (Magura group of nappes,
Czech Republic). Geol. Carpathica 50, 1, 33—48.

Bukry D. 1973: Low-latitude coccolith biostratigraphic zonation.

Init. Rep. DSDP 15, 685—703.

Burnett J.A. 1990: A new nannofossil zonation scheme for the Bo-

real Campanian. INA Newsletter 12, 67—70.

Burnett J.A. 1997: New species and new combinations of Cretaceous

nannofossils, and a note on the origin of Petrarhabdus (Deflan-
dre) Wind & Wise. J. Nannoplankton Res. 19, 2, 133—146.

Burnett J.A. 1998: Upper Cretaceous. In: Bown P.R. (Ed.): Calcar-

eous Nannofossil Biostratigraphy. Cambridge University
Press, 132—199.

Burnett J.A. & Whitham F. 1999: Correlation between the nanno-

fossil and macrofossil biostratigraphies and the lithostratigra-
phy of the Upper Cretaceous of NE England. Proc. Yorkshire
Geol. Soc. 52, 4, 371—381.

Heck S.E.v. & Prins B. 1987: Refined Nannoplankton Zonation for

the Danian of the Central North Sea. In: Stradner H. & Perch-
Nielsen K. (Eds.): International Nannoplankton Association,
Vienna Meeting 1985, Proceeding. Abh. Geol. Bundesanst.
39, 285—303.

Hofmann T., Švábenická L. & Hradecká L. 1999: Biostratigraphy

of Fissure Fillings in the Ernstbrunn Limestone of the Wasch-
berg Zone (Lower Austria). Abh. Geol. B.—A. 56/2, 465—474.

Kaever M. 1980: Geologie des westlichen Münsterlandes. Führer zu

vor- und frühgeschichtlichen Denkmälern, Mainz, 45, 20—39.

Kaever M. & Lommerzheim A. 1995: The highest Campanian in

the transitional area between the boreal and the Tethyan
realms in NW-Germany: Implications for stratigraphy, facies,
palaeoceanography and palaeoclimatology. Münstersche For-
sch. Geol. Paläont. 77, 293—327.

Lommerzheim A.J. 1995: Stratigraphie und Ammonitenfaunen des

Santons und Campans im Münsterländer Becken (NW-Deut-
schland). Geol. Paläont. Westf. 40, 3—97.

Malata T. & Poprawa P. 1997: Subsidence and uplift analysis of

the Polish part of Outer Carpathian basins – backstripping of
reconstructed profiles of the basin-fill. In: Krobicki M. &
Zuchiewicz W. (Eds.): Dynamics of the Pannonian-Car-
pathian-Dinaride System. Przegl. Geol. 45, 10, 1088—1089.

Matějka A. & Roth Z. 1948: Report of map sheet Vsetín. Věst.

Stát. Geol. Úst. 23, 133—136 (in Czech).

Perch-Nielsen K. 1985: Mesozoic calcareous nannofossils. In:

Bolli H.M., Saunders J.B. & Perch-Nielsen K. (Eds.): Plank-
ton Stratigraphy. Cambridge University Press, 329—426.

Pesl V. 1968: Lithofacies of Paleogene in the Magura Unit of the

Outer Carpathian Flysch on the territory of Czechoslovakia
and Poland. Sbor. Geol. Vied, Rad ZK 9, 71—117 (in Czech,
English summary).

Pospichal J.J. & Wise W.W. 1990: Maastrichtian calcareous nan-

nofossil biostratigraphy of maud Rise ODP Leg 113 Sites 689
and 690, Weddell Sea. Proc. ODP, Sci. Results 113, 465—487.

Potfaj M. 1993: Position and role of the Biele Karpaty Unit in the

Flysch Zone of the West Carpathians. Geol. Práce, Spr. 98,
55—78 (in Slovak, English summary).

Potfaj M. 1998: Geodynamics of the Klippen Belt and Flysch Belt

of the Western Carpathians. In: Rakús M. (Ed.): Geodynamic
development of the Western Carpathians. Geol. Survey of Slo-
vak Republic, 143—154.

Sissingh W. 1977: Biostratigraphy of Cretaceous nannoplankton,

with appendix by Prins B. & Sissingh W. Geol. en Mijnb. 56,
37—65.

Stráník Z., Krejčí O. & Menčík E. 1989: Contribution to the geolo-

gy of the Biele Karpaty Unit. Zpr. Geol. Výzk. v Roce 1986
143—147 (in Czech).

Stráník Z., Bubík M., Čech S. & Švábenická L. 1996: The Upper

Cretaceous in South Moravia. Věst. Čes. Geol. Úst. 71, 1, 1—30.

Summesberger H., Švábenická L., Čech S., Hradecká L. & Hof-

mann  T.  1999:  New  palaeontological  and  biostratigraphical
data  on  the  Klement  and  Pálava  Formations  (Upper  Creta-
ceous)  in  Austria  (Waschberg-Ždánice  Unit).  Ann.  Naturhist.
Mus. Wien 100A, 39—79.

Švábenická L. 1992: Upper Cretaceous nannofossils from the Kle-

ment Formation (Flysch Belt of the West Carpathians, Czecho-
slovakia). Proceedings of the 4

INA Conf., Knih. ZPN 14a,

189—205.

Švábenická L. 1995: The stratigraphical corellation of the Campa-

nian low- and high-latitude calcareous nannofossils in South-
ern Moravia (Western Carpathians). Geol. Carpathica 46,
297—302.

Švábenická L. 1998: Occurrence of Petrarhabdus copulatus (De-

flandre) Wind and Wise in the Outer Western Carpathians.
Věst. Čes. Geol. Úst. 73, 2, 89—98.

Švábenická L. 1999: Penetration of high-latitude nannoflora to the

depositional area of the Outer Western Carpathians in the Turo-
nian-Maastrichtian. Geol. Carpathica 50, Spec. Issue, 77—79.

Švábenická L. & Bubík M. 1992: The fossil record (calcareous

nannofossils and foraminifers) in single intervals of the Up-
per Cretaceous flysch sequence of the West Carpathians,
Czechoslovakia. Cretaceous Research 13, 583—590.

Švábenická L., Bubík M., Krejčí O. & Stráník Z. 1997: Stratigra-

phy of Cretaceous sediments of the Magura group of nappes
in Moravia. Geol. Carpathica 48, 3, 179—191.

Švábenická L., Wagreich M. & Egger H. in press: Late Cretaceous

nannofossil assemblages at a transect from the northern Tethys
to the temperate realm in Central Europe. In: Michalík J. (Ed.):
Tethyan/Boreal Cretaceous Correlation. Veda, Bratislava.

Thierstein H. 1976: Mesozoic calcareous nannoplankton biostratig-

raphy of marine sediments. Mar. Micropaleont. 1, 325—362.

Thierstein H.R. 1981: Late Cretaceous nannoplankton and the

change at the Cretaceous-Tertiary boundary. Soc. Econ. Pale-
ont. Min. Spec. Publ. 32, 355—394.

Thierstein H. & Haq B.U. 1977: Maastrichtian/Danian biogeo-

graphic variations in calcareous nannoplankton (Abstr.). J.
Paleontology 51, Suppl., 28—29.

Tomek Č. 1996: PANCARDI – Dynamics of Ongoing Orogeny.

In: Gee D.G. & Zeyen H.J. (Eds.): EUROPROBE 1996 –
Lithosphere Dynamics: Origin and Evolution of Continents.
Uppsala University, 15—26.

Watkins D.K. 1992: Upper Cretaceous nannofossils from Leg 120,

Kerguelen Plateau, Southern Ocean. Proc. ODP, Sci. Results
120, 343—370.

CAMPANIAN/MAASTRICHTIAN PENETRATION OF HIGH-LATITUDE NANNOFLORA 39

Watkins D.K., Wise S.W., Pospichal J.J. & Crux J. 1996: Upper

Cretaceous  calcareous  nannofossil  biostratigraphy  and  pale-
oceanography  of  the  Southern  Ocean.  In:  Moguilevsky  A.  &
Whatley  R.  (Eds.):  Microfossils  and  Oceanic  Environments.
University of Wales, Aberystwyth Press, 355—381.

Wind F.H. 1979: Maastrichtian-Campanian nannofloral provinces

of the Southern Atlantic and Indian oceans. In: Talwani M. &
Hay W. et al. (Eds.): Deep Sea Drilling Results in the Atlantic
Ocean: Continental Margins and Paleoenvironment. Amer.
Geophys. Union 123—127.

Wise S.W. Jr. 1993: Mesozoic and Cenozoic calcareous nannofos-

sils recovered by Deep Sea Drilling Project Leg 71 in the
Falkland Plateau region, Southwest Ocean. Init. Rep., Deep
Sea Drilling Project 71, 481—550.

Worsley T.R. 1974: The Cretaceous-Tertiary boundary event in the

ocean. SEPM Spec. Publ. 20, 94—125.

Worsley T.R. & Martini E. 1970: Late Maastrichtian nannoplank-

ton provinces. Nature 225, 1242—1243.

Appendix 1

Nannofossil taxa identified at the localities mentioned in text:

1.1 – Hluk (Púchov Marl, tectonic slice in the front of the Bílé Karpaty
Unit, Magura group of nappes); Late Campanian—?basal Maastrichtian,
zone UC15d

—UC16.

Ahmuellerella  octoradiata,  A.  regularis,  Arkhangelskiella  cymbiform-
is,  Biscutum  coronum,  B.  ellipticum,  Broinsonia  enormis,  Broinsonia
parca  constricta,  B.  p.  parca,  Ceratolithoides  aculeus,  Cretarhabdus
conicus,  Cribrosphaerella  ehrenbergii,  Cyclagelosphaera  sp.,  Cylin-
dralithus  serratus,  Eiffellithus  gorkae,  E.  parallelus  (one  badly  pre-
served  specimen),  E.  turriseiffelii,  Gartnerago  obliquum,  Kamptneri-
us  magnificus,  Lithraphidites  carniolensis,  Manivitella  pemmatoidea,
Markalius inversus, Microrhabdulus attenuatus, M. decoratus, Micula
staurophora,  M.  swastica,  Petrarhabdus  copulatus,  Placozygus  fibuli-
formis, Prediscosphaera cretacea, P. grandis, P. majungae, P. ponticu-
la, P. spinosa, P. stoveri, Quadrum gartneri, Reinhardtites levis, Reta-
capsa  angustiforata,  R.  schizobrachiata,  Rotelapillus  crenulatus,
Rucinolithus  wisei,  Staurolithites  mielnicensis,  Tranolithus  phacelo-
sus,  Uniplanarius  gothicus,  U.  sissinghii,  U.  trifidus,  Watznaueria
barnesae,  W.  biporta,  Zeugrhabdothus  bicrescenticus,  Z.  diplogram-
mus, Z. embergeri, Z. sigmoides, curved spines (sensu Burnett 1997, p.
145. Plate 1: Fig. 27).

1.2 – Javořina 42D

/95 (Javorina Formation, Bílé Karpaty Unit, Magu-

ra group of nappes); Early/Late Campanian boundary, zone UC15c

Arkhangelskiella  cymbiformis,  Biscutum  melaniae,  Braarudosphaera
bigelowii,  Broinsonia  parca  constricta,  Ceratolithoides  aculeus,  Cre-
tarhabdus  conicus,  Cribrosphaerella  ehrenbergii,  Cyclagelosphaera
sp.,  Eiffellithus  eximius,  E.  gorkae,  E.  pospichalii,  Eprolithus  floralis,
Glaukolithus  compactus,  Kamptnerius  magnificus,  Manivitella  pem-
matoidea,  Micula  staurophora,  Microrhabdulus  attenuatus,  M.  deco-
ratus,  Petrarhabdus  copulatus,  Placozygus  cf.  sigmoides,  Predis-
cosphaera  cretacea,  P.  grandis  (fragments),  P.  spinosa,  Reinhardtites
anthophorus,  Retacapsa  crenulata,  Thoracosphaera  sp.,  Tranolithus
phacelosus, Uniplanarius sissinghii, Watznaueria barnesae, W. bipor-
ta, W. quadriradiata, Zeugrhabdotus embergeri.

1.3  –  Racková  60B  (Soláň  Formation,  Rača  Unit,  Magura  group  of
nappes); Late Maastrichtian, zone UC20.
Acuturris  scotus,  Ahmuellerella  octoradiata,  A.  regularis,  Arkhangel-
skiella cymbiformis, A.  cf. maastrichtiana,  Biantholithus sparsus, Bis-
cutum  coronum,  Braarudosphaera  bigelowii,  Chiastozygus  antiquus,
C.  litterarius,  Corollithion  exiguum,  C.  ?madagaskarensis,  Cri-
brosphaerella  ehrenbergii,  Eiffellithus  gorkae,  E.  parallelus,  E.
pospichalii,  Gartnerago  obliquum,  Helicolithus  trabeculatus,  Kampt-
nerius  magnificus,  Lithraphidites  carniolensis,  L.  praequadratus-

quadratus,  L.  quadratus,  Lucianorhabdus  inflatus,  Micula  stauropho-
ra,  Microrhabdulus  attenuatus,  M.  decoratus,  Neocrepidolithus  cohe-
nii,  Octolithus  multiplus,  Placozygus  fibuliformis,  P.  sigmoides,  Pre-
discosphaera arkhangelskyi, P. cretacea, P. grandis, P. cf. majungae,
P.  spinosa,  P.  stoveri,  Psyktosphaera  cf.  firthii,  Retacapsa  crenulata,
Rhagodiscus angustus, R. asper, Staurolithites mielnicensis, Tegumen-
tum  stradneri,  Watznaueria  barnesae,  Zeugrhabdotus  bicrescenticus,
Z. embergerii, Z. cf. trivectis.

1.4 – Valašské Meziříčí (Soláň Formation, Rača Unit, Magura group
of nappes); upper part of Early Campanian, zone UC15b

Arkhangelskiella cymbiformis, A. specillata, Biscutum coronum, B. el-
lipticum, B. melaniae, Broinsonia enormis, Calculites obscurus, Cera-
tolithoides  aculeus,  Chiastozygus  litterarius,  Cretarhabdus  conicus,
Cribrosphaerella  ehrenbergii,  Cyclagelosphaera  sp.,  Eiffellithus  exi-
mius,  E.  gorkae,  Gartnerago  obliquum,  Kamptnerius  magnificus,  K.
punctatus,  Lithastrinus  grillii,  Lucianorhabdus  ex  gr.  cayeuxii,  L.  in-
flatus,  L.  maleformis,  Manivitella  pemmatoidea,  Markalius  inversus,
Microrhabdulus  decoratus,  Micula  staurophora,  Micula  swastica,
Neocrepidolithus  watkinsii,  Placozygus  fibuliformis,  Prediscosphaera
cretacea,  P.  grandis  (fragments),  P.  ponticula,  P.  spinosa,  ?Quadrum
svabenickae, Reinhardtites anthophorus, R. cf. R. levis, Retacapsa an-
gustiforata,  R.  schizobrachiata,  Rucinolithus  sp.,  Staurolithites  cf.  S.
mielnicensis,

Tranolithus

phacelosus,

Uniplanarius

gothicus,

Watznaueria barnesae, W. biporta, W. britannica, Zeugrhabdotus bi-
crescenticus, Z. embergeri, Z. praesigmoides.

Appendix 2

Nannofossil taxa mentioned in the text, in alphabetical order of genera
epithets.

Acuturris scotus (Risatti 1973) Wind & Wise in Wise & Wind 1977
Ahmuellerella octoradiata  (Górka  1957)  Reinhardt  1966
Ahmuellerella regularis (Górka 1957) Reinhardt & Górka 1967
Amphizygus brooksii Bukry 1969
Angulofenestrelithus  snyderi  Bukry  1969
Arkhangelskiella  confusa  Burnett  1998
Arkhangelskiella cymbiformis Vekshina 1959
Arkhangelskiella  maastrichtiana  Burnett  1998
Arkhangelskiella specillata  Vekshina  1959
Biantholithus sparsus Bramlette & Martini 1964
Biscutum coronum Wind & Wise in Wise & Wind 1977
Biscutum dissimilis Wind & Wise in Wise & Wind 1977
Biscutum ellipticum (Górka 1957) Grün & Alleman 1975
Biscutum magnum Wind & Wise in Wise & Wind 1977
Biscutum melaniae (Górka 1957) Burnett 1997
Biscutum notacalum Wind & Wise in Wise & Wind 1977
Braarudosphaera bigelowii (Gran & Braarud 1935) Deflandre 1947
Braarudosphaera  turbinea  Stradner  1963
Broinsonia enormis (Shumenko 1968) Manivit 1971
Broinsonia matalosa (Stover 1966) Burnett in Gale et al. 1996
Broinsonia parca constricta Hattner et al. 1980
Broinsonia parca parca (Stradner 1963) Bukry 1969
Broinsonia signata (Noël 1969) Noël 1970
?Bukrylithus  ambiguus  Black  1971
Calculites obscurus (Deflandre 1959) Prins & Sissingh in Sissingh 1977
Calculites ovalis (Stradner 1963) Prins & Sissingh in Sissingh 1977
Ceratolithoides  aculeus  (Stradner  1961)  Prins  &  Sissingh  in  Sissingh

1977

Ceratolithoides arcuatus Prins & Sissingh in Sissingh 1977
Ceratolithoides  kamptneri  Bramlette  &  Martini  1964
Chiastozygus  antiquus  (Perch-Nielsen  1973)  Burnett  1998
Chiastozygus litterarius  (Górka  1957)  Manivit  1971
Corollithion  exiguum  Stradner  1961
Corollithion  ?madagaskarensis  Perch-Nielsen  1973
Corollithion  signum  Stradner  1963
Cretarhabdus conicus Bramlette & Martini 1964

40 ŠVÁBENICKÁ

Cribrocorona gallica  (Stradner  1963)  Perch-Nielsen  1973
Cribrosphaerella  daniae  Perch-Nielsen  1973
Cribrosphaerella ehrenbergii (Arkhangelsky 1912) Deflandre in Piveteau 1952
Cruciellipsis  cuvillieri  (Manivit  1966)  Thierstein  1971
Cyclagelosphaera reinhardtii  (Perch-Nielsen  1968)  Romein  1977
Cylindralithus  biarcus  Bukry  1969
Cylindralithus serratus Bramlette & Martini 1964
Dodekapodorhabdus  noeliae  Perch-Nielsen  1968
Eiffellithus eximius (Stover 1966) Perch-Nielsen 1968
Eiffellithus  gorkae  Reinhardt  1965
Eiffellithus  parallelus  Perch-Nielsen  1973
Eiffellithus  pospichalii  Burnett  1998
Eiffellithus turriseiffelii (Deflandre in Deflandre & Fert 1954) Reinhardt 1965
Eprolithus moratus (Stover 1966) Burnett 1998
Eprolithus floralis (Stradner 1962) Stover 1966
Gartnerago obliquum (Stradner 1963) Noël 1970
Grantarhabdus coronadventis (Reinhardt 1966) Grün in Grün & Alle-

mann 1975

Haqius circumradiatus (Stover 1966) Roth 1978
Helicolithus trabeculatus (Górka 1957) Verbeek 1977
Helicolithus turonicus Varol & Girgis 1994
Heteromarginatus bugensis (Górka 1957) Crux in Crux et al. 1982
Kamptnerius  magnificus  Deflandre  1959
Kamptnerius  punctatus  Stradner  1963
Lapideacassis  cf.  L.  bispinosa  (Perch-Nielsen  in  Perch-Nielsen  &

Franz 1977) Burnett 1998

Lithastrinus  grillii  Stradner  1962
Lithastrinus  septenarius  Forchheimer  1972
Lithraphidites  carniolensis  Deflandre  1963
Lithraphidites  praequadratus  Roth  1978
Lithraphidites  quadratus  Bramlette  &  Martini  1964
Lucianorhabdus  cayeuxii  Deflandre  1959
Lucianorhabdus  inflatus  Perch-Nielsen  &  Feinberg  in  Perch-Nielsen

1986

Lucianorhabdus maleformis Reinhardt 1966
Manivitella pemmatoidea (Deflandre in Manivit 1965) Thierstein

1971

Markalius apertus Perch-Nielsen 1979
Markalius inversus (Deflandre in Deflandre & Fert 1954) Bramlette &

Martini 1964

Marthasterites furcatus (Deflandre in Deflandre & Fert 1954) Deflandre

1959

Microrhabdulus  attenuatus  (Deflandre  1959)  Deflandre  1963
Microrhabdulus  decoratus  Deflandre  1959
Micula concava (Stradner in Martini & Stradner 1960) Verbeek 1976
Micula murus (Martini 1961) Bukry 1973
Micula  prinsii  Perch-Nielsen  1979
Micula staurophora (Gardet 1955) Stradner 1963
Micula swastica Stradner & Steinmetz 1984
Misceomarginatus pleniporus Wind & Wise in Wise & Wind 1977
Monomarginatus pectinatus Wind & Wise in Wise & Wind 1977
Monomarginatus quaternarius Wind & Wise in Wise & Wind 1977
Nannoconus  sp.
Neocrepidolithus  cohenii  (Perch-Nielsen  1968)  Perch-Nielsen  1984
Neocrepidolithus watkinsii Pospichal & Wise 1990
Nephrolithus frequens Górka 1957
Octocyclus reinhardtii (Bukry 1969) Wind & Wise in Wise & Wind 1977
Octolithus multiplus  (Perch-Nielsen  1973)  Romein  1979
Orastrum  campanensis  (Cepek  1970)  Wind  &  Wise  in  Wise  &  Wind

1977

Petrarhabdus copulatus (Deflandre 1959) Wind & Wise in Wise 1983
Placozygus fibuliformis  (Reinhardt  1964)  Hoffmann  1970
Podorhabdus?  elkefensis  Perch-Nielsen  1981
Prediscosphaera  arkhangelskyi  (Reinhardt  1965)  Perch-Nielsen  1984
Prediscosphaera cretacea (Arkhangelsky 1912) Gartner 1968
Prediscosphaera  grandis  Perch-Nielsen  1979
Prediscosphaera  majungae  Perch-Nielsen  1973
Prediscosphaera ponticula  (Bukry  1969)  Perch-Nielsen  1984
Prediscosphaera spinosa (Bramlette & Martini 1964) Gartner 1968
Prediscosphaera stoveri (Perch-Nielsen 1968) Shafik & Stradner 1971
Psyktosphaera firthii Pospichal & Wise 1990
Quadrum gartneri Prins & Perch-Nielsen in Manivit et al. 1977
Quadrum  svabenickae  Burnett  1998
Reinhardtites  anthophorus  (Deflandre  1959)  Perch-Nielsen  1968
Reinhardtites levis Prins & Sissingh in Sissingh 1977
Retacapsa  angustiforata  Black  1971
Retacapsa  crenulata  (Bramlette  &  Martini  1964)  Grün  in  Grün  &  Allemann

1975

Retacapsa ficula (Stover 1966) Burnett 1998
Retacapsa schizobrachiata (Gartner 1968) Grün in Grün & Allemann

1975

Rhagodiscus angustus (Stradner 1963) Reinhardt 1971
Rhagodiscus asper (Stradner 1963) Reinhardt 1967
Rhagodiscus infinitus (Worsley 1971) Applegate et al. in Covington &

Wise 1987

Rotelapillus crenulatus (Stover 1966) Perch-Nielsen 1984
Rucinolithus ? magnus Bukry 1975
Rucinolithus wisei Thierstein 1971
Seribiscutum primitivum (Thierstein 1974) Filewicz et al. in Wise &

Wind 1977

Sollasites horticus (Stradner et al. in Stradner & Adamiker 1966) Ce-

pek & Hay 1969

Staurolithites  aachenus  (Bukry  1969)  Burnett  1998
Staurolithites  ellipticus  (Gartner  1968)  Lambert  1987
Staurolithites  imbricatus  (Gartner  1968)  Burnett  1998
Staurolithites  laffittei  Caratini  1963
Staurolithites  mielnicensis  (Górka  1957)  Perch-Nielsen  1968
Thoracosphaera operculata Bramlette & Martini 1964
Tortolithus hallii (Bukry 1969) Crux in Crux et al. 1982
Tranolithus gabalus Stover 1966
Tranolithus minimus (Bukry 1969) Perch-Nielsen 1984
Tranolithus orionatus  (Reinhardt  1966)  Reinhardt  1966
Tranolithus phacelosus Stover 1966
Uniplanarius gothicus (Deflandre 1959) Hattner & Wise 1980
Uniplanarius  sissinghii  Perch-Nielsen  1986
Uniplanarius  trifidus  (Stradner  in  Stradner  &  Papp  1961)  Hattner  &

Wise 1980

Watznaueria barnesae (Black 1959) Perch-Nielsen 1968
Watznaueria biporta Bukry 1969
Watznaueria britannica (Stradner 1963) Reinhardt 1964
Watznaueria fossacincta Wind & Cepek 1979
Watznaueria quadriradiata Bukry 1969
Zeugrhabdotus bicrescenticus (Stover 1966) Burnett in Gale et al.

1996

Zeugrhabdotus diplogrammus (Deflandre in Deflandre & Fert 1954)

Burnett in Gale et al. 1996

Zeugrhabdotus embergeri (Noël 1958) Perch-Nielsen 1984
Zeugrhabdotus praesigmoides Burnett 1998
Zeugrhabdotus sisyphus (Gartner 1968) Crux 1989
Zeugrhabdotus trivectis Bergen 1994