GEOLOGICA CARPATHICA, 51, 4, BRATISLAVA, AUGUST 2000
CALCAREOUS DINOFLAGELLATE AND CALPIONELLID
BIOEVENTS VERSUS SEA-LEVEL FLUCTUATIONS RECORDED
IN THE WEST-CARPATHIAN (LATE JURASSIC/EARLY CRETACEOUS)
Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, 842 26 Bratislava, Slovak Republic; firstname.lastname@example.org
(Manuscript received September 28, 1999; accepted in revised form March 15, 2000)
Abstract: Recently established separate dinoflagellate cyst zonation combined with the successive calpionellid events
contribute to the HIRES of the Upper Jurassic and Lower Cretaceous Tethyan pelagic carbonate sequences. Composi-
tional changes in dinoflagellate and calpionellid assemblages are correlated with eustatic sea-level fluctuations. Thus,
parallel calpionellid and cyst zonations give us more precise tools for the subdivision of deposits investigated as well
as for better understanding and reconstruction of the paleoceanographical and paleoecological conditions of the an-
cient marine environments. The calcareous resting cyst distribution is shown to be influenced by the whole complex
of environmental factors such as sea-level transgressive/regressive pulses, hydrological regime, nutrient content etc.
Key words: Upper Jurassic, Lower Cretaceous, Western Carpathians, calcareous dinoflagellates, calpionellids, integrated
biochronology, paleoecology, sea-level changes.
Review of previous calcareous dinoflagellate studies
Plankton has an important role in the ecology of the ocean.
Besides calpionellids, there were calcareous dinoflagellates
which represented a further substantional planktonic ele-
ment during the Late Jurassic and Early Cretaceous. Since
Kaufmanns description (in Heer 1865) of Lagena ovalis
and Lagena sphaerica, many notions of single chambered
bodies of various shape and size (between 4063
m in di-
ameter) appeared in the literature. They were attributed to
newly defined genera Stomiosphaera Wanner 1940, Cadosi-
na Wanner 1940, Pithonella Lorenz 1902, and Calcisphaer-
ula Bonet 1956. The shape, size, gross structure and optical
properties of the tests sectioned were the main taxonomical
criteria for their classification. Contemporaneously, some
authors (Colom 1955; Wanner 1940; Vogler 1941; Bonet
1956; Durand Delga 1957; Leischner 1959; Nagy 1966,
1971; Lineckaya 1974; etc.) pointed out that the vertical dis-
tribution of dinoflagellate associations may be used for time
correlation of the pelagic sequences.
A new stage in these studies started after Bollis (1974)
publication including documentation of Jurassic and Creta-
ceous calcisphaeres isolated from soft Indian Ocean sedi-
ments in the framework of the Deep Sea Drilling Project.
Scanning electron microscope (SEM) observations were
used in investigation of specimens studied. Bolli (1974) de-
scribed nineteen new species and four forms in open no-
menclature of Pithonella Lorenz 1902 and included them in
the incertae sedis family Calcisphaerulidae. However, he
entirely omitted taxonomical diagnoses of the previously
established above mentioned families. Bolli (l.c.) supposed
that comparative studies of the topotypic material including
optical and SEM methods might be realized in the future.
On the other hand, he hoped that his newly established spe-
cies would not be synonymous with previously registered
ones. However, the fact emerged, that several species suc-
cessfully used in the biostratigraphic zonation during the
last 40 years (Nowak 1966, 1968, 1976; Borza 1980a, 1984;
Borza & Michalík 1986) were classified in the genus Pitho-
nella. Later, (Øehánek 1992; Øehánek & Heliasz 1993;
Vaíèek et al. 1994; Lakova et al. 1999; Reháková 2000)
solved some new problems of cyst biozonation.
Wall & Dale (1968) for the first time pointed out the possi-
bility of the genetic interpretation of calcisphaeres as calcar-
eous dinoflagellate cysts. Taking into consideration this fact,
as well as the results of Fütterer (1976), Keupp (1981, 1987)
showed the same nature for the Mesozoic calcisphaerulids.
Keupp (1987) established the calcareous dinoflagellate sys-
tem based on the orientation of the calcite crystals forming
the outer calcareous wall layer. He subdivided all known cyst
genera of the order Peridiniales Haeckel 1894 into three sub-
families: Orthopithonelloideae, Obliquipithonelloideae and
Pithonelloideae. Thus, the taxonomy of calcareous di-
noflagellate cysts became more approximate to the biological
classification. This approach was dynamically followed by
Keupps group (Keupp & Mutterlose 1984; Keupp 1980,
1984, 1987, 1990, 1991, 1992; Fütterer 1990; Willems 1988,
1990, 1992, 1994; Keupp & Ilg 1989; Keupp & Versteegh
1989; Keupp & Kowalski 1992; Keupp et al. 1992; etc.).
In an accordance with Keups division, Øehánek (in Øe-
hánek & Cecca 1993) compared cadosinids and stomio-
sphaerids with dinoflagellate cysts on the basis of the optical
character of their sections in polarized light. He included
genera Stomiosphaera Wanner 1940, Colomisphaera Nowak
1968, Committosphaera Øehánek 1985, Parastomiosphaera
Nowak 1968, Carpistomiosphaera Nowak 1968 and Stomi-
osphaerina Nowak 1974 into the subfamily Orthopitonel-
loideae Keupp 1987. On the other hand, genera Cadosina
Wanner and Crustocadosina Øehánek 1985 belong to the
subfamily Obliquipithonelloideae Keupp 1987. A different
opinion was presented by Colom (1994) who summarized
accessible knowledge on dinoflagellate cysts from the Bale-
aric Islands area. He compared the small spherical, sporadi-
cally conical forms of the incertae sedis group with the re-
cent genera Gromia, Allogromia, Difflugia, Trigonopyris.
All forms presented were included by him in the Ca-
dosinidae, Stomiosphaeridae and Calcisphaerulidae families.
A new contribution to systematic concepts was presented by
Reháková & Michalík (1996), who used both the combined
optical and SEM methods on Early Cretaceous cyst speci-
mens. They proved that the wall structure of Cadosina fusca
Wanner is identical to Obliquipithonella multistrata (Pflau-
mann & Krashenninikov 1978), the type species of the
Obliquipithonelloideae Keupp 1987. They obtained the same
results in Stomiosphaera wanneri Borza 1969 which ought to
be identical with Orthopithonella congruens Fütterer 1990, a
typical representative of the Orthopithonelloideae Keupp
1987. According to Hildebrand-Habel & Willems (1997),
these investigations should be acceptable in a new approach
to the systematics of calcareous dinoflagellates.
The main goal of this paper is to show the calcareous di-
noflagellates distribution as a tool for more detailed bios-
tratigraphy of carbonate pelagic sequences as well as for the
interpretation of the paleoenvironmental conditions. This
work also takes into account the tendency of the Committee
of Stratigraphy and Paleontology of the Carpathian-Balkan
Geological Association to create an integrated biostrati-
graphical scale, practically acceptable for the Upper Juras-
sic and Lower Cretaceous sedimentary records of the whole
The first calcareous cyst zonal scheme (including 6 Late
Kimmeridgian to Hauterivian cyst zones) was proposed by
Nowak (1968) who later on, (Nowak 1976) revised it. Fur-
ther contributions to the biostratigraphy of calcisphaerids
were made by Borza (1969, 1984), Borza & Michalík
(1986), Øehánek (1992), Øehánek & Heliasz (1993), Øe-
hánek & Cecca (1993), Vaíèek et al. 1994. Lakova et al.
(1999) not only confirmed all previously proposed dinocyst
zones in the West Balkan and West Fore-Balkan area, but
also recognized a further three Upper Berriasian to Valang-
inian dinocyst interval-zones.
The last detailed biostratigraphical contribution was made
by Reháková (2000), who investigated the vertical distribu-
tion of the calcareous dinoflagellates in Late Oxfordian to
Upper Albian sedimentary sequences of the Western Car-
pathians. On the basis of a series of successive first occur-
rences and acme accumulations of calcareous dinoflagellates
she proposed a separate cyst biozonation. In this paper, re-
corded dinocyst events are directly correlated to the calpi-
onellid ones (sensu Reháková & Michalík 1997a), to the am-
monite zonation (Hoedemaeker et al. 1993; Cariou &
Hantzpergue 1997), to the stratigraphic time scale (Gradstein
et al. 1995) as well as to the sea-level fluctuation (Haq et al.
1988, Fig. 1). This approach offers good arguments for estab-
lishing an integrated high-resolution event stratigraphic
(HIRES) scale of the West-Carpathian Upper Jurassic and
Lower Cretaceous pelagic sequences which will also be sup-
ported by radiolarian, planktonic foraminiferal and nanno-
plankton distribution in a short time. The sections and forma-
tions studied, their geological position in the framework of
the West-Carpathian tectonic units are described in Vaíèek
et al. (1994), Reháková (1995).
Oxfordian stage of the calcareous dinocyst evolution
Lower Oxfordian passage beds are characterized by the
first occurrence (FO) of Cadosina parvula Nagy (Pl. I:
Figs.12) which indicates the base of the Parvula Zone (Re-
háková 2000). Fragments of dysaerobic bivalves (Oschmann
1995) form a persistent, substantional part of the Oxfordian
microfacies (Pl. I: Fig. 8). Shortly after the FO of Colo-
misphaera fibrata (Nagy) Pl. I: Figs. 34, the acme of this
index species was documented in the uppermost Oxfordian
deposits (Reháková 2000). The acme of C. fibrata coincides
with the onset of a sea-level rise (Haq et al. 1988). The as-
semblage also contains rare Colomisphaera pieniniensis
(Borza) Pl. I: Fig. 5 and Schizosphaerella minutissima
(Colom) Pl. I: Fig. 6. A rich calcareous dinoflagellate as-
sociation of the same age was documented by Keupp & Ilg
(1989) from the shallower coastal parts of Normandy. At the
end of the Oxfordian the abundance of bivalves was decreas-
ing. In a well-oxygenated setting planktonic foraminifers be-
came prevailing (Mutterlose & Böckel 1998). Hauslerina
helvetojurassica (Hausler) Pl. I: Fig. 7 and Globuligerina
bathoniana (Pazdrowa) are present in the West-Carpathian
pelagic sediments. In order to show their rock-forming role,
pre-Kimmeridgian Protoglobigerinae Zone was distin-
guished by Dragastan et al. (1975) in the East-Carpathian re-
gion. The Favusellacea became holoplanktonic (fully plank-
tonic) and according to Simmons et al. (in BouDagher-Fadel
et al. 1997) their sudden appearance may have been related
to a rising eustatic sea-level, which opened up new niches.
Kimmeridgian stage of cyst evolution
Cadosina parvula became most abundant at the beginning of
the Kimmeridgian. On the basis of this event, the Parvula
Acme Zone was distinguished (Reháková 2000). The index spe-
cies is accompanied by common cysts of Schizosphaerella
minutissima (Colom) and Colomisphaera carpathica (Borza)
Pl. I: Fig. 9. Planktonic foraminifers have been still domi-
nating in microfacies of this time. Mass abundance of Globo-
Fig. 1. The most important calpionellid and cyst bioevents in the
West-Carpathian area corelated with the combined ammonite zona-
tion (Hoedemaeker et al. 1993; Gradstein et al. 1995) and eustatic
sea-level fluctuations sensu Haq et al. (1988).
DINOFLAGELLATE AND CALPIONELLID BIOEVENTS VS. SEA-LEVEL FLUCTUATIONS 231
Plate I: Fig. 1. Cadosina parvula Nagy. Oxfordian, Czorsztyn Unit, bar = 100
m. Fig. 2. Abundant Cadosina parvula Nagy. Early Kim-
meridgian, Czorsztyn Unit, bar = 200
m. Figs. 34. Colomisphaera fibrata (Nagy). Late Oxfordian, Vysoká Unit, bars = 50
m. Fig. 5.
Colomisphaera pieniniensis (Borza). Late Oxfordian, Czorsztyn Unit, bar = 50
m. Fig. 6. Schizosphaerella minutissima (Colom). Kim-
meridgian, Vysoká Unit, bar = 50
m. Fig. 7. Hauslerina helvetojurassica (Hausler). Late Oxfordian, Czorsztyn Unit, bar = 50
8. Microfacies with abundant juvenile bivalve fragments. Oxfordian, Manín Unit, bar = 200
m. Fig. 9. Colomisphaera carpathica (Bor-
za). Kimmeridgian, Pieniny Unit, bar = 50
DINOFLAGELLATE AND CALPIONELLID BIOEVENTS VS. SEA-LEVEL FLUCTUATIONS 233
Plate II: Fig. 1. Microfacies with Globochaete alpina Lombard. Kimmeridgian, Manín Unit, bar = 100
m. Fig. 2. Saccocoma packstone.
Kimmeridgian, Czorsztyn Unit, bar = 200
m. Fig. 3. Colomisphaera nagyi (Borza). Kimmeridgian, Czorsztyn Unit, bar = 50
m. Fig. 4.
Stomiosphaera moluccana Wanner. Kimmeridgian, Czorsztyn Unit, bar = 50
m. Fig. 5. Carpistomiosphaera borzai (Nagy). Late Kim-
meridgian, Vysoká Unit, bar = 50
m. Fig. 6. Colomisphaera pulla (Borza). Early Tithonian, Vysoká Unit, bar = 50
m. Fig. 7. Carpistomi-
osphaera tithonica Nowak. Early Tithonian, Vysoká Unit, bar = 50
m. Fig. 8. Parastomiosphaera malmica (Borza). Early Tithonian,
Czorsztyn Unit, bar = 100
m. Fig. 9. Microfacies with acme C. pulla. Early Tithonian, Czorsztyn Unit, bar = 100
m. Fig. 10. Microfacies
with Parastomiosphaera malmica (Borza). Early Tithonian, Pruské Unit, bar = 100
chaete alpina Lombard (Pl. II: Fig. 1) proves favourable en-
vironmental conditions for the development of green algae.
There was also the acme of Saccocoma Agassiz (Pl. II: Fig.
2). Planktonic crinoids became rock-forming organisms.
This fact led Dragastan et al. (1975) to define the Saccocoma
Zone. According to Matyszkiewicz (1997) and Keupp &
Matyszkiewicz (1997) saccocomids are abundant in facies
which prograded on the epicontinental platforms of the pas-
sive northern Tethyan shelf during the Late Oxfordian/?Ear-
liest Kimmeridgian and Late Kimmeridgian/Early Tithonian
and they marks the late transgressive systems tract as well as
the presumed high stand deposits. Calcareous dinoflagellate
associations were also abundant and diversified. Among
them, groups with orthopitonellid type of wall structure re-
flecting pelagic conditions (Mutterlose & Böckel 1998) were
dominant. Colomisphaera nagyi (Borza) Pl. II: Fig. 3 oc-
curs rarely in the higher part of the Kimmeridgian sequences
together with Stomiosphaera moluccana Wanner (Pl. II: Fig.
4), an index species of the Moluccana Zone (sensu Nowak
1976). Carpistomiosphaera borzai (Nagy) Pl. II: Fig. 5)
appears in the uppermost Kimmeridgian deposits. Nowak
(l.c.) defined the top Kimmeridgianlowermost Tithonian
Borzai Zone, later accepted by Borza (1984). According to
Reháková (2000), the Borzai Zone is considered to be of
Late Kimmeridgian age only.
Tithonian dinocyst stage
The dinocyst form Colomisphaera pulla (Borza) Pl. II:
Fig. 6 appeared at the beginning of the Early Tithonian.
Nowak (1968) used the FO of this form for definition of the
Pulla Zone, which he later abandoned. Borza (1984) pointed
out a synchroneous FO event of Colomisphaera pulla and
Carpistomiosphaera tithonica Nowak (Pl. II: Fig. 7) on the
basis of which he proposed his Pulla-Tithonica Zone. De-
tailed studies of several sections indicated that the interval
with abundant C. pulla (Pl. II: Fig. 9) precedes the FO of C.
tithonica. This ecoevent was regarded as the Pulla Acme
Zone (Reháková 2000). Environmental conditions dominat-
ing during the Early Tithonian were very favourable for de-
velopment of calcareous dinocyst associations. Abundant
Parastomiosphaera malmica (Borza) Pl. II: Figs. 8, 10
appeared in the upper part of Lower Tithonian deposits.
Carpistomiosphaera tithonica was also a distinct form of this
interval. The FO of these two species was used for defining
another two dinocyst Tithonica and Malmica zones. All the
above mentioned Lower Tithonian cyst zones are character-
ized by high abundance and high diversity of dinoflagellate
associations and they coincide with an elevated eustatic sea-
level (Fig. 1).
The Middle Tithonian (sensu Gradstein et al. 1995) di-
nocyst zones Colomisphaera minutissima and C. carpathica
were distinguished by Nowak (1968). Later, on the basis of
the FO of Colomisphaera cieszynica Nowak, this interval
was redefined as the Cieszynica Zone (Nowak 1976). Be-
cause, the first microgranular calpionellid forms of the genus
Chitinoidella were shown to be a more reliable tool in strati-
fication of pelagic deposits, these above mentioned cyst
zones were not accepted (Borza 1984). However, the recent
tendency leading to utilization of the widest spectrum of ele-
ments suitable for detailed subdivision has forced the spe-
cialists to create separate dinoflagellate zonation. Correlating
Kimmeridgian to Tithonian dinoflagellate and ammonite as-
sociations from the Monte Nerone pelagic limestone in Italy,
Øehánek (in Øehánek & Cecca 1993) re-established the
Cieszynica Zone. However, his conclusions have not been
confirmed by Reháková (2000) who found out that the FO of
Cadosina semiradiata semiradiata Wanner (Pl. III: Figs.12)
precedes the FO of chitinoidellids belonging to the Dobeni
Subzone. On the basis of these facts, the Semiradiata di-
nocyst zone has been distinguished (Reháková 2000).
The FO of Colomisphaera tenuis (Nagy) Pl. III: Figs.
34, an index species of the Tenuis Zone, was also docu-
mented in the Middle Tithonian part of the pelagic sequence.
It coincides with the onset of more advanced and diversified
chitinoidellids of the Boneti Subzone. In the uppermost part
of the Middle Tithonian deposits Colomisphaera fortis Øe-
hánek (Pl. III: Figs. 56) appeared. This index species was
used for defining of the Fortis Zone by Øehánek (1992). This
zone was later accepted by Lakova et al. (1999). It is also ac-
ceptable in the West-Carpathian sequence. It is worth men-
tioning, that the interval with C. fortis contains a dinoflagel-
late association poor in both, abundance and diversity. It
coincides with an abrupt ecoevent in the calpionellid associa-
tion: chitinoidellid exctinction. Later, an ecological change
may have been triggered their substitution by hyaline calpi-
onellid forms of the Praentintinnopsella and the Crassicollar-
ia zones. Shortly before the first occurrence of hyaline calpi-
onellids, marks of distinct erosion and redeposition with
sedimentary breccia layers was observed in several of the
sections studied. The overlying strata usually contain no
chitinoidellids, but the first transitional calpionellids have an
inner hyaline and an outer microgranular wall layer. Breccia
layers from the Chitinoidella and Praetintinnopsella transi-
tion beds are known practically from the whole West-Car-
pathian area. Although they were studied in detail from the
Vysoká Unit of the Krína Nappe (Reháková & Michalík
1995), they were not named. It seems, that the global third-
order sea-level fall, called here the Hlboè Event (Fig. 1),
was also associated with a rapid turnover in calpionelid evo-
lution. Afterwards, favourable environmental conditions for
the development of calpionellids predominated. Among
qualitatively new hyaline associations, several radiations,
stagnant and extinction phases were documented. Shortly af-
ter the FO of Tintinnopsella remanei Borza, the index species
of the Remanei Subzone, abundant crassicollarian forms of
larger size (Crassicollaria intermedia (Durand Delga) and
Cr. massutiniana (Colom)) appeared. This radiation coin-
cides with the third-order sea-level rise as is shown in Fig. 1.
The FO of abundant small crassicollarian forms (Cr. brevis
Remane) characterize the Brevis Subzone of the Crassicol-
laria Zone. Shortly after their appearance, large crassicollari-
ans disappeared. This stagnant calpionellid phase coincides
with the third-order sea-level fall. Calpionella grandalpina
Nagy, C. alpina Lorenz and Crassicollaria parvula Remane
dominated in a new radiation phase. The occurrence of abun-
dant Schizosphaerella minutissima is observed as a coeval di-
DINOFLAGELLATE AND CALPIONELLID BIOEVENTS VS. SEA-LEVEL FLUCTUATIONS 235
Plate III: Figs. 12. Cadosina semiradiata semiradiata Wanner. Middle Tithonian, Manín Unit, bars=50
m. Figs. 34. Colomisphaera
tenuis (Nagy). Middle Tithonian, Czorsztyn Unit, bars=50
m. Figs. 56. Colomisphaera fortis Øehánek. Middle Tithonian, Vysoká Unit,
m. Figs. 78. Stomiosphaerina proxima Øehánek. Late Tithonian, Vysoká Unit, bars=50
m. Fig. 9. Microfacies with Cadosina
semiradiata fusca Wanner and Tintinnopsella carpathica (Murg. & Filip.). Berriasian, Czorsztyn Unit, bar=50
m. Figs. 1012. Deformed
(?aberrant) crassicollarians. Late Tithonian, Zliechov Unit, bars=100
m. Fig. 13. Favusella hoterivica (Subbotina). Late Berriasian, Pruské
m. Figs. 1415. Gonoglobuligerina gulekhensis (Gorbachik, Poroshina). Early Valanginian, Pruské Unit, bars=100
nocyst event coinciding with an interval of calpionellid max-
imum diversity and it can be correlated with the sea-level
Decrease in the abundance of calcareous dinoflagellates is
documented in the Upper Tithonian sequence. The FO of rare
Stomiosphaerina proxima Øehánek (Pl. III: Figs. 78) was
identified. The rare cadosinids (Cadosina semiradiata fusca
(Wanner) and C. semiradiata semiradiata (Wanner)) are also
present in microfacies of this time. According to Øehánek
(1992), the FO of the index species S. proxima (defining the
Proxima cyst zone) characterizes the Jurassic-Cretaceous
boundary in the Western Carpathians. The Proxima Zone is
accepted by Reháková (2000), although its age is regarded as
Late Tithonian. The same result was arrived at by Lakova et
There is also an extinction of highly diversified crassicol-
larians which happened across the Tithonian-Berriasian
boundary (Pl. III: Fig. 10). Compared with the interval of
chitinoidellid disappearance, the same scenario of the envi-
ronmental behaviour was also documented during the inter-
val of crassicollarian retreat. Marks of erosion accompanied
by siliclastic input and breccia accumulations were identified
accross the whole West-Carpathian area (Pl. IV: Fig. 1).
Huge, several metres thick breccia bodies observed in the
Zliechov Unit of the Krína Nappe (Michalík et al. 1995) can
serve as a suitable example of environmental turnover. This
abrupt change of the sedimentary conditions in the West-Car-
pathian area is defined as the Zliechov Event. It coincides
with a global third-order sea-level fall (Fig. 1) interpreted as
the so-called Purbeckian regression (Zakharov et al. in
Rawson et al. 1996). Shortly afterwards, abundant aberrant
crassicollarian forms (Pl. III: Figs. 1112) occurred in envi-
ronments influenced by a distinct siliclastic input. More tur-
biditic water masses and enhanced productivity could lead to
diminishing penetration of sunlight into the photic zone (Gil-
bert & Clark 19821983). These conditions were not optimal
for calcareous dinoflagellates and calpionellids.
On a global scale, the Jurassic Cretaceous Boundary Bio-
Event is characterized as a second-order mass extinction in-
terval. According to Barnes et al. (1996) it was spread
through three short-term extinction events, or steps, at the
base, middle and the end of the Tithonian Stage. Three dis-
tinct extinction steps are also documented among the plank-
tonic associations of this time: saccocomid extinction during
the Early Tithonian, chitinoidellid extinction during the Mid-
dle Tithonian and crassicollarian extinction during the Late
BerriasianValanginian dinocyst stage
Development of the calcareous dinoflagellates persisted in
its stagnant phase from the Middle Berriasian. Rare
Schizosphaerella minutissima and Stomiosphaerina proxima
only were observed in sedimentary sequences of this time in-
terval. Beside the dominant nannoconid associations, envi-
ronmental conditions were favourable for calpionellid devel-
opment. Free niches opened by the crassicollarian extinction
were occupied by the expanding (r-strategist) spherical
Calpionella alpina Lorenz, an index species of the Alpina
Subzone of the standard Calpionella Zone (Allemann et al.
1971; Remane et al. 1986). This form created a nearly mono-
specific association, which persisted from the appearence of
the first remaniellids indicating the Ferasini Subzone. After a
certain time following the innovation, strong calpionellid di-
versification is observable from the uppermost part of the El-
liptica Subzone to the middle part of the standard Calpionel-
lopsis Zone (the Oblonga Subzone). Abundant dinocyst
(Cadosina semiradiata fusca, Pl. III: Fig. 9) with an
obliquipithonelloid structure of calcite crystals forming a
double-layered wall (Reháková & Michalík 1996) appeared
in the interval of increasing calpionellid diversity. An inter-
val with accumulation of this index species was considered
as the Fusca Acme Zone (Reháková 2000). The abundance of
these cysts varies from 3550 % on pelagic elevations to 6
10 % of the planktonic remnants in basinal bottom sedi-
ments. After a longer break lasting from the end of Early
Kimmeridgian, planktonic foraminifers represented by Fa-
vusella hoterivica (Subbotina) Pl. III: Fig. 13 and Gono-
globuligerina gulekhensis (Gorbachik & Poroshina) Pl.
III: Figs. 14, 15 appeared in the planktonic assemblage. En-
hanced calcareous dinoflagellate and calpionellid produc-
tion, as well as the sudden onset of non-keeled, globular for-
aminifers, organisms typical of the Boreal bioprovince
(Gasinski 1997), coincide with a second-order eustatic rise
(Reháková & Michalík 1997b). It seems that a similar short
communication between the biota of adjacent provinces as
was documented during the Late Oxfordian, was repeatedly
renewed during the Late Berriasian sea-level highstand.
An onset of more pelagic facies accompanied by both, dis-
tinct change in micro- and macrofaunal composition as well
as changes recorded in stable isotopic values and clay miner-
als were discussed by Adatte et al. (1996). Sea-level rise in-
fluenced the atmospheric and consequently also the hydrody-
namic oceanic regime. Near to the Calpionella and
Calpionellopsis zonal boundary frequent intercalations of ra-
diolaria rich horizons (Pl. IV: Fig. 2) appear in the hitherto
rather monotonous calpionellid wackestones indicating more
intensive aeration of deeper layers of oceanic water influ-
enced by upwelling activity Reháková (1998).
A tiny dinocyst form, Stomiosphaera wanneri Borza (Pl.
IV: Figs. 34) with typical orthopithonelloid wall structure
(Reháková & Michalík 1996) was identified in the upper-
most part of the Upper Berriasian. The FO of this index spe-
cies is considered to be the base of the Wanneri cyst zone
which was established by Lakova et al. (1999). Their results
have also been confirmed by Reháková (1999). The Wanneri
Zone corresponds to the Late Berriasian calpionellid Oblon-
ga and Murgeanui subzones of the Calpionellopsis Zone. At
the end of this zone a distinct breccia accumulation known as
the Nozdrovice Breccia (Borza et al. 1980b; Pl. IV: Fig. 5)
was identified in several sections. The third-order eustatic
curve shows a rapid fall of the sea-level (Fig. 1). Shortly af-
terwards, sudden siliclastic input disturbing previously mo-
notonous basinal carbonate sedimentation was observed.
This broadly identified environmental change (similar to those
shown in Late Tithonian) negatively influenced the amount of
microplankton components. Calcareous dinoflagellates de-
DINOFLAGELLATE AND CALPIONELLID BIOEVENTS VS. SEA-LEVEL FLUCTUATIONS 237
Plate IV: Fig. 1. Brecciated limestone from the Jurassic-Cretaceous boundary interval representing the global sea-level fall (named as the
Zliechov Event). Tatric Superunit, bar = 200
m. Fig. 2. Radiolarian wackestone reflecting the Late Berriasian current regime change. Kysu-
ca Unit, bar = 200
m. Figs. 34. Stomiosphaera wanneri Borza. Late Berriasian, Vysoká Unit, bars = 50
m. Fig. 5. Nozdrovice Breccia
with Calpionella alpina Lorenz. Late Berriasian, Zliechov Unit, bar = 100
m. Figs. 67. Aberrant calpionellid forms documented in the
end of standard Calpionellopsis Zone. Late Berriasian, Zliechov Unit, bars = 50
m. Fig. 8. Acme Minuta (microfacies with abundant Cado-
sina minuta Borza). Early Valanginian, Czorsztyn Unit, bar = 100
creased in abundance. Previously highly diversified calpionel-
lid associations rapidly decreased in diversity and abundance,
too. Abundant aberrant calpionellid forms were observed in
many of the studied sections (Pl. IV: Figs. 6, 7). This regres-
sive pre-phase, leading later to calpionellid extinction, ulti-
mately caused an increase in the evolutionary rate of nanno-
From the topmost part of the Lower Valanginian deposits
the FO of the cyst form of Cadosina minuta Borza (Pl. IV: Fig.
8) was confirmed. On the basis of the abundant monoassocia-
tion of the index species, this distinct short interval was de-
fined as the Minuta Acme Zone (Reháková 2000). Among a
nannoconid blooming the calpionellids tried to survive. Only
several large forms of the standard Calpionellites Zone (Dard-
eri and Major subzones) successfully asserted themselves in a
strong selection stress. A very brief new radiation calpionellid
phase coincides with a small third-order sea-level rise on a
broad second-order sea-level fall. The Upper Valanginian se-
quence contains the dinocyst form Colomisphaera vogleri
(Borza) Pl. V: Figs. 12. A little later, the FO of Carpisto-
miosphaera valanginiana Borza (Pl. V: Figs. 34) was ob-
served. The appearance of these two index species were used
for establishing the Vogleri and Valanginiana cyst zones. Dif-
ferent results were obtained by Ivanova (in Lakova et al. 1999)
who stated that the FO of the above mentioned index species
are synchronous and they together characterize the onset of the
Carpistomiosphaera valanginiana Zone.
A new, stronger siliclastic input (Pl. V: Fig. 5) represented
by the Oravice Event coinciding with the rapid third-order
sea-level fall (Fig. 1). This abrupt change in environmental
conditions led to total calpionellid decimation in almost the
whole Tethyan region. It seems that this was the reason why
calpionellids have never been observed in the Boreal Realm.
On the other hand, rapid evolution and spreading of nanno-
conid communities is documented as a coeval event to the
calpionellid crisis (Pl. V: Fig. 6). Only rare Tintinnopsella
carpathica (Murgeanu & Filipescu) survived in the huge
nannoconid blooms until the Late Valanginian, where marly
limestones show a short interval of nannoconid depletion.
From that point, overlying thin turbiditic intercalations con-
tain rich accumulations of bivalve fragments (Pl. V: Fig. 7)
recording coeval low oxygenate conditions in the adjacent
areas. The marks of widespread-levels Late Valanginian
transgression (Mutterlose 1992) controlled by further envi-
ronmental factors were recorded from the Outer West-Car-
pathian area (Michalík et al. 1995). According to their inter-
pretation, a positive excursion of the
C corresponded to a
short warm and humid climate interval preceding the mid-
Cretaceous greenhouse state. Locally graded intercalations,
rich in radiolaria and sponges (Pl. V: Fig. 8) could have
been linked with the periodically active contour currents
persisting until the Early Hauterivian.
Hauterivian Barremian dinocyst stage
The cyst form Stomiosphaera echinata Nowak (Pl. V:
Figs. 910) was documented in the Upper ValanginianLate
Barremian passage beds. The FO of this rare form was stated
as the base of the Echinata Zone. This interval contains a low
abundance, but a highly diversified dinocyst association. A
coeval more diversified cyst association from the NW part of
the German Basin was described by Keupp (1979, 1981).
The maximum diversity of calcareous dinoflagellates coin-
cides with the trangressive phase recorded by the second-or-
der eustatic curve (Fig. 1). An ongoing transgression influ-
enced a turbiditic regime documented practically throughout
the whole West-Carpathian area. The huge Stráovce turbid-
ite complex (Pl. V: Fig. 11) deposited in the Zliechov Basin
of the Fatric Unit was described by Borza et al. (1980b). The
factor responsible for its accumulation is regarded here as the
Stráovce Event. The evident depletion in dinoflagellate di-
versification is observed in Lower Barremian deposits. Nev-
ertheless, the nannofloral speciation and development have
Aptian dinocyst stage
The disappearance of the index species forms of Colo-
misphaera vogleri and Stomiosphaera echinata at the begin-
ning of the Early Aptian was considered as the base of the
Cieszynica-Olzae cyst zone (Reháková 2000), in which ca-
dosinids became dominant in the dinocyst association. The
zone was named according to synchronously appearing
obliquipithonelloid indexes: Cadosina semiradiata cieszyni-
ca (Nowak) Pl. VI. Fig. 1 and C. semiradiata olzae
(Nowak) Pl. VI: Fig. 2. Only one orthopithonelloid dino-
form Colomisphaera heliosphaera (Vogler) survived. The
Early Aptian global climatic change (Arthur et al. 1991;
Weissert & Lini 1991) is mirrored in the basinal Carpathian
environments (Michalík et al. 1999), too. An onset of the
black shale deposition was documented in several of the stud-
ied sections. The most spectacular occurrence from the Kysu-
ca Basin was described as the Koòhora Event which was cor-
related with the known Selli Event of Erba (1994).
Halásová in (Michalík et al. 1999) parallelized a dramatic de-
crease in abundance of the Nannoconus with the event known
as the nannoconid crisis (Erba 1994). Planktonic foramini-
fers became dominant components of planktonic communities.
Marly limestone with rich accumulations of radiolarians and
sponges, periodically intercalated by black shale sequence,
point to a renewed contourite current activity.
At the beginning of the Middle Aptian new forms of mi-
crogranular calpionellids appeared in foraminiferal wacke-
stones to packstones. The vertical span of the calpionellid
genera Praecolomiella Borza, Deflandronella Trejo, and
Parachitinoidella Trejo was defined as the Praecolomiella
Zone. Microgranular praecolomiellids are less frequent, but,
on the other hand, their loricas are twice or several times
larger than those of Middle Tithonian chitinoidellids. The no-
mismogenesis of planktonic foraminifers lowered the selec-
tional stress among the calpionellids and this competitive en-
vironment led to a growth of their loricas. The revival of
microgranular calpionellids at this time level allows us to
speculate about a similar climatic and paleoceanographic
conditions as were previously described in the Middle Titho-
nian (see position of the eustatic curve on Fig. 1).
DINOFLAGELLATE AND CALPIONELLID BIOEVENTS VS. SEA-LEVEL FLUCTUATIONS 239
Plate V: Figs. 12. Colomisphaera vogleri (Borza). Late Valanginian, Vysoká Unit, bars = 50
m. Figs. 34. Carpistomiosphaera valangini-
ana Borza. Late Valanginian, Manín Unit, bars = 50
m. Fig. 5. Sandy limestone deposited during the Oravice Event. Early Valanginian,
Zliechov Unit, bar = 100
m. Fig. 6. Nannoconid packstone. Late Valanginian, Czorsztyn Unit, bar = 100
m. Fig. 7. Calcareous turbidite
limestone with bivalve fragments. Late Valanginian, Pruské Unit, bar = 100
m. Fig. 8. Sponge packstone documented in the Lower Hau-
terivian part of Pruské Unit, bar = 100
m. Figs. 910. Stomiosphaera echinata Nowak. Early Hauterivian, Czorsztyn Unit, bars = 50
Fig. 11. Pelbiodetritic limestone with benthonic foraminiferal remnants sedimented in turbidite regime during the Stráovce Event. Hauteriv-
ian, Zliechov Unit, bar = 100
Plate VI: Fig. 1. Cadosina semiradiata cieszynica (Nowak). Early Aptian, Manín Unit, bar=50
m. Fig. 2. Cadosina semiradiata olzae
(Nowak). Early Aptian, Manín Unit, bar=50
m. Fig. 3. Cadosina oraviensis Borza. Late Albian, Tatric Superunit, bar=50
m. Fig. 4. Microfa-
cies with Calcisphaerula innominata Bonet. Late Albian, Campanian congl. from the Pieniny Klippen Belt (PKB) area, bar=100
m. Fig. 5.
Stomiosphaera sphaerica Bonet. Late Albian, Tatric Superunit, bar=50
m. Fig. 6. Microfacies with Pithonella ovalis (Kaufmann). Late Albi-
an, Manín Unit, bar=100
m. Fig. 7. Colomisphaera gigantea (Borza). Late Albian, Tatric Superunit, bar=50
m. Fig. 8. Microfacies with
abundant planktonic foraminifers. Albian, Kysuca Unit, bar=100
m. Fig. 9. Microfacies with Pithonella trejoi Bonet, Bonetocardiella
conoidea (Bonet), Calcisphaerula innominata Bonet and Stomiosphaera sphaerica Bonet. Late Albian, Ilerdian congl. of the PKB, bar=100
DINOFLAGELLATE AND CALPIONELLID BIOEVENTS VS. SEA-LEVEL FLUCTUATIONS 241
Albian dinocyst stage
Something like a restriction phase of calcareous dinocyst
production is observable at the beginning of the Albian. On
the other hand, there was a new explosive phase of nanno-
conid evolution documented by Erba & Quadrio (1987). Mi-
crogranular calpionellid forms disappeared. They were sub-
stituted by a new group with hyaline loricas. The FO of these
hyaline calpionellids is used for definition of the last Early
Cretaceous calpionellid Colomiella Zone which includes all
species of the Colomiella Bonet and Calpionellopsella Trejo.
It seems that microgranular calpionellid forms gave rise to
hyaline ones several times independently. The change of the
lorica composition was synchroneous with the Early Albian
peak in nannoconid abundance (Erba & Quadrio 1987) simi-
larly, as during the previously described Late Tithonian
change of a chitinoidellid microgranular structure (Reháková
& Michalík 1997a). The development of the last two men-
tioned calpionellid associations coincided with the elevated
rate of the third-order sea-level rise (Fig. 1).
From the Middle to the Late Albian, there were very favour-
able environmental conditions for calcareous dinoflagellate
development in the West-Carpathian area. Their innovation
and radiation phases can be correlated with a broad second-or-
der eustatic rise (Fig. 1). Two dinocyst zones, Cadosina ora-
viensis and Calcisphaerula were previously distinguished by
Borza in (Borza & Michalík 1986). According to Reháková
(2000), the interval with abundant Cadosina oraviensis Borza
(Pl. VI: Fig. 3) is considered as the Oraviensis Acme Zone.
The index species is further accompanied by abundant Cadosi-
na callosa Knauer usually occurring in sediments character-
ized by foraminiferal and crinoidal microfacies (Pl. VI: Fig.
8). The Late Albian interval with abundant Calcisphaerula in-
nominata Bonet (Pl. VI: Fig. 4) was considered as the Innomi-
nata Acme Zone (Reháková 2000). Shortly above the FO of
the index species, Stomiosphaera sphaerica (Kaufman) Pl.
VI: Fig. 5, Pithonella ovalis (Kaufmann) Pl. VI: Fig. 6, Co-
lomisphaera gigantea (Borza) Pl. VI: Fig. 7, Bonetocardi-
ella conoidea (Bonet) and Pithonella trejoi Bonet (Pl. VI: Fig.
9) appear in the glauconitic limestone and marly sequence.
Dinoflagellates formed a significant element of the Juras-
sic and Cretaceous marine phytoplankton throughout the
world in open shelf, slope and basinal environments. Due to
very favourable conditions for the development of plankton-
ic associations, a rich and structured ecosystems could orig-
inate in the photic zone of the Tethyan Realm during that
time. Certain dinoflagellate taxa formed a resistant calcare-
ous/or sporopollenin cyst which was the only potentionally
fossilizable stage of their life cycle. The focus of this study
was precisely calcareous dinocyst associations. The investi-
gation of their vertical distribution allowed us:
a) to show that the independent dinocyst zonation can
serve as one of the important tools of the integrated bios-
tratigraphy of the Upper Jurassic and Lower Cretaceous car-
bonate deposits of the Western Carpathians;
b) to correlate this zonation with dinocyst zonations estab-
lished recently in the East-Carpathian area, in order to show
its interregional availability;
c) to distinguish several diversification and diversity re-
duction events among the cyst associations studied and to
utilize them for paleoenvironmental reconstruction.
d) to correlate the dinocyst zonation with the calpionellid
events and zonation and thus to contribute to the HIRES of
the Upper Jurassic and Lower Cretaceous Tethyan pelagic
It seems that not only calpionellids but also calcareous di-
noflagellates belonged to the planktonic elements sensitive-
ly recording the whole complex of environmental changes
such as climate perturbations, sea-level fluctuations, nutri-
ent distribution. It was shown that the sea-level transgres-
sive stages were favourable for dinocyst development and
all distinguished acme concentrations of cyst taxa studied
were controlled by sea-level highstand phases. On the other
hand, cyst diversity reduction events coincided with sea-
level regressive stages.
Acknowledgements: The paper was prepared in the frame-
work of the Grant Project GAV No. 2/6058/99. Author ex-
press her sincere thanks to Dr. I. Lakova, Prof. J. Remane,
Prof. Dr. H. Keupp, Assoc. Prof. J. Michalík and to Prof. M.
Miík, for their critical reading of manuscript, their com-
ments, and for profitable discussion on Jurassic-Cretaceous
paleoecology and paleoceanography. The photographs were
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