GEOLOGICA CARPATHICA, FEBRUARY 2008, 59, 1, 71—85
The Central Paratethys was a large intracontinental sea
consisting of a chain of sedimentary basins extending
through the Alpine-Carpathian region. These basins were
periodically connected with the Meditteranean Sea, to
the Indopacific and to the Atlantic Oceans, and some-
times isolated (Rögl & Steininger 1983; Rögl 1998,
1999; Fig. 1).
Foraminiferal species diversity is considered here as the
number of foraminiferal species in a certain time interval.
The analysis of species diversity changes in the Central
Paratethys has been based on the assumption that the evo-
lution of species diversity is influenced by different fac-
tors in the open ocean and in the inland basin due to peri-
ods of connections and isolations.
The analysed time interval includes the period from
the Late Eocene to the Sarmatian/Pannonian boundary
(37—11.6 Ma; Berggren et al. 1995; Cicha et al. 1998;
Gradstein et al. 2004).
The goal of the contribution presented here is to define
intervals with marked changes of species diversity and to
suggest their causes.
Foraminiferal species diversity in the Central Paratethys
– a reflection of global or local events?
Department of Geology and Paleontology, Charles University in Prague, Albertov 6, CZ-128 43 Praha 2, Czech Republic;
(Manuscript received March 6, 2006; accepted in revised form October 10, 2007)
Abstract: The stratigraphical ranges of the Eocene—Sarmatian Central Paratethys Foraminifera were used to calculate
their species diversity ( = the number of foraminiferal species in a certain time interval). Species diversity was evaluated
separately for benthic and planktonic Foraminifera, agglutinated and porcellanaceous foraminifers, lagenids, oxyphilic,
dysoxic and euryhaline species. In the Central Paratethys basins, six foraminiferal species diversity events were defined:
(i) The Egerian species diversity maximum was reached by progressive growth in the number of foraminiferal species
during approximately 15 Myr from the Early Eocene to the Egerian. (ii) The Egerian and Egerian/Eggenburgian
extinction event interrupted the growth of species diversity. The beginning of extinction was approximately isochronous
with the acme of small-sized reticulofenestras (calcareous nannoplankton) which are considered to be an indicator of
warm water with low nutrient concentrations; (iii) The Ottnangian species diversity minimum is the consequence of a
minimum number of foraminiferal FOs during the Eggenburgian and the Ottnangian (3 Myr). This was caused by
isolation of the Paratethys and the progressive decrease of salinity and oxygen content during the Ottnangian. (iv) The
Early Badenian species diversity maximum followed an abrupt increase in species diversity of benthic Foraminifera at the
Karpatian/Badenian boundary as a consequence of a large transgression and the establishment of fully marine conditions
in nearly the whole territory. Diversity of the planktonic assemblages increased gradually in the Karpatian (1.1
a result of the opening of new sea-ways. (v) Extinction at the Badenian/Sarmatian boundary indicates rapid shift of
paleoenvironmental parameters. (vi) The number of Sarmatian species represents only 29 % of the Egerian species. The
composition of assemblages changed and shifted to endemic, euryhaline species. These changes can be clearly correlated
with the isolation and vanishing of the Central Paratethys basins. It is suggested that foraminiferal species diversity in the
intracontinental Central Paratethys basins was controlled mainly by regional paleogeographic communication/isolation
events which may be influenced by global sea-level changes and partly by regional paleoenvironmental changes
influenced by global climate changes.
Key words: Oligocene, Miocene, Central Paratethys, paleoenvironment, paleogeography, species diversity, Foraminifera.
1.1 Global events in the world oceans
Three fundamental parameters triggered global changes
in the world oceans with consequent species diversity
variations: (1) astronomical variation in the Earth’s orbit
and inclination, namely the Milankovitsh Cycles (Hays et
al. 1976; Lourens & Hilgen 1997; Zachos et al. 2001)
(23,000; 41,000 and 95,000 years periodicity); (2) fluctua-
tions in the production of oceanic crust: when rates of pro-
duction of oceanic crust decrease, the sea-level falls and
decreases, land area increases and epi-
continental seas decreases, which may lead to climatic
cooling (Berger et al. 1981; Barron 1985); (3) extraterres-
trial bodies impact events (Alvarez et al. 1980; Coccioni
et al. 2000).
For the analysed time interval (37—11.6 Ma), the follow-
ing global events were defined: (1) the terminal Eocene
event characterized by marked cooling, decreased latitudi-
nal thermal gradients and a gradual decrease in the fertility
of the ocean related to a major reorganization of the water
masses. The influence of extraterrestrial events (microtec-
tite-rich layer) may have accelerated some processes
(Keller 1986; Pomerol & Premoli-Silva 1986; Berggren &
Prothero 1992; Prothero 1994; Spezzaferri et al. 2002a;
Prothero et al. 2003).
(2) The general cooling observed in the Miocene and
Pliocene were defined as Mi-events (Miller et al. 1991; Turco
et al. 2001; Billups & Schrag 2002). In the studied interval
between 37—11.6 Ma, the following Mi-events were identi-
fied: Mi1 event – 23.8 Ma, Mi1a event – 22 Ma, Mi1b
event – 17.5 Ma, Mi2 event – 16.5 Ma, Mi3 event –
13.8 Ma, Mi4 event – 13.2 Ma, Mi5 event – 11.7 Ma. A
good correlation was observed between the sea-level curve
of Haq et al. (1988) and Mi-events (Billups & Schrag 2002).
In the Central Paratethys the 3rd-order global cycles TA
4.4—4.5, TB.1.1—1.5 and TB.2.1—2.6 can be correlated to se-
quences from the studied time interval (Kováč et al. 2001).
1.2 Foraminiferal species diversity in the world oceans
In the analysed time interval, Kaiho (1994) defined two
major global foraminiferal extinction events: the first in
Late Eocene and the second in the early Middle Miocene
affecting both surface- and deep-water species.
The Late Eocene event occurred during 38—34 Ma
(Boersma 1986; Keller et al. 1992; Spezzaferri et al.
2002a). The extinction percentages were calculated as
67 % for planktonic foraminifers predominantly surface
dwelling species, 14 % for intermediate-water species and
9—46 % for deep-dwelling species (Kaiho 1994). The
cause of the Eocene extinction is interpreted as a step by
step cooling of low-latitude surface water.
The early Middle Miocene extinction interval was dat-
ed between 17—13 Ma (Thomas 1986; Thomas & Vincent
1987; Berggren & Miller 1989). Extinction percentages
were calculated as 40 % for planktonic species and 35—
52 % for benthic intermediate- and deep-water species.
The causes of this event have been explained by water
mass changes resulting from warming (associated with
sluggish circulation) and subsequent cooling in high-lat-
itude surface and deep-water (Thomas & Vincent 1987;
Woodruff & Savin 1991).
In addition, moderate low-oxygen events during the
Late Oligocene (27—25 Ma) were documented by Kaiho
(1991). These events were without dramatic extinctions
except for deep-water benthic foraminifers. The event co-
incided with an increase in ocean-crust production, which
increased the sea level (Keigwin & Keller 1984), and the
warm-water species among planktonic foraminifers (Hal-
lock et al. 1991).
1.3 Foraminiferal diversity in the Central Paratethys
Species diversity in periodically isolated land-locked
marine basins is controlled by both global and regional
events (= regional tectonic and communication/isolation
events). In comparison with the oceanic realm, the Central
Paratethys consisted of small shallow bodies characterized
by higher spatial and temporal variability.
In the real oceanic realm the only parameters influenc-
ing foraminiferal distribution are: amount of organic mat-
ter (food supply), oxygen concentration both in water and
sediment, temperature, salinity and sedimentological char-
acteristics (Corrlis 1981; Sjoerdsma & Van der Zwaan
1992; Jorrisen et al. 1992; Gooday 1994; Van der Zwaan
et al. 1999; Haslett 2002; Smart 2002). However in an in-
land sea, a much broader range of variation is expected in
the same environmental parameters, even to the extreme.
The analysis of foraminiferal species diversity from the
Central Paratethys was based on stratigraphical ranges of
species summarized in the atlas “Oligocene, Miocene For-
aminifera of the Central Paratethys” (Cicha et al. 1998) for
the interval from the Upper Eocene to the Sarmatian. In
this atlas, stratigraphical ranges were given for every spe-
cies in relation to the following stratigraphical intervals:
Late Eocene, early Kiscellian, late Kiscellian, Egerian,
Eggenburgian, Ottnangian, Karpatian, Badenian (divided
into the substages Moravian, Wielickian and Kosovian)
and Sarmatian. The atlas synthesizes data obtained during
long-term work by great numbers of co-authors. At present
the atlas represents the most suitable input data for analy-
sis of foraminiferal species diversity although some inac-
curacies in stratigraphical distributions of species are ex-
pected. These errors may also influence evaluation of
species diversity changes. Piller in Harzhauser & Piller
(2007) used the same data pool (Cicha et al. 1998) for def-
inition of extinction and immigration/origination events
during the Miocene but he selected only 425 species-level
taxa for event definition. Biostratigraphical events (Last
Occurrence (LO) and First Occurrence (FO)) were defined
at a stage boundary or within individual stages and sub-
stages. This type of data enables the calculation of the fol-
lowing characteristics for individual stratigraphical inter-
vals (stages or substages):
(i) Total numbers of foraminiferal species were counted
separately for (1) benthic species, (2) planktonic species,
(3) the main taxonomic groups: Textulariina, Miliolina
and Lagenina, (4) groups with specific paleoecological re-
quirements: dysoxic ( = Bolivina spp., non-costate Bulimi-
na spp., Uvigerina spp., Fursenkoina spp.; Sen-Gupta &
Machain-Castillo 1993; Loubere 1994; Kaiho 1994,
1999; Bernhard et al. 1997; Den Dulk et al. 1998, 2000;
Spezzaferri et al. 2002b; Báldi 2006), oxyphilic (= cibici-
doids, Lenticulina spp., miliolids based on data from
Kaiho 1999; Den Dulk et al. 2000; Spezzaferri et al.
2002b; Báldi 2006) and euryhaline ( = elphidiids, miliol-
Fig. 1. Sketch of the Paratethys and its connection with surround-
ing basins (from Rögl & Steininger 1983).
FORAMINIFERAL SPECIES DIVERSITY IN THE CENTRAL PARATETHYS
ids, Ammonia div. sp.; some agglutinated foraminifers
(Ammobaculites spp., Eggerella spp.); Phleger 1960; Mur-
ray 1973, 1991).
The relative numbers (percentages) of these groups in
benthic foraminiferal assemblages were also calculated.
(ii) The numbers of FOs and LOs in the stages (substages)
and at the stage (substage) boundaries were calculated sepa-
rately for all groups enumerated in the previous paragraph.
(iii) Extinction percentages (Kaiho 1994) and corre-
sponding “appearance percentages” were calculated for
stages and stage boundaries based on the formulas given
below. In the Central Paratethys basins, this index quanti-
fies regional appearance/disappearance events.
EP = * 100 [%] AP = * 100 [%]
EP – local extinction percentage calculated for stage or
– number of LOs at the upper stage
– number of LOs in the stage, AP – local
appearence percentage calculated for stage or substage,
number of FOs at the lower stage boundary, N
number of FOs in the stage or substage, N
– number of
foraminiferal species in stage or substage.
(iv) The numbers of bioevents (LOs and FOs) in individ-
ual stages and substages were standardized.
BE/1 Myr =
BE/1 Myr – standardized value of numbers of bioev-
– number of LOs in the stratigraphical unit,
– number of FOs in the stratigraphical unit, d
ration of the stratigraphical unit (range of stratigraphical
units from Steininger et al. 1990; Cicha et al. 1998; Kováč
2000; Gradstein et al. 2004; Hohenegger et al. 2007).
It was expected that the species diversity in a certain
stratigraphical interval was determined by (i) global envi-
ronmental parameters of this interval (e.g. sea-level low- or
highstand, cold/warm period, oxygen concentration, os-
cillations of salinity, global nutrient level, etc.) and (ii)
facial diversifications: species diversity of foraminifers
reaches higher values in facially more diversified basins.
In relation to isolation/communication events, the fol-
lowing foraminiferal species may form assemblages: (1)
taxa surviving from the previous interval, (2) new immi-
grants from neighbouring seas or oceans indicating com-
munication, (3) new endemic species which may indicate
3.1 Changes of foraminiferal species diversity
3.1.1 Benthic species (Fig. 2)
The values for species diversity show two maximum
peaks (Fig. 2A) and vary from 71 species in the Sarmatian
to 247 species in the Egerian and 231 in the Moravian. Di-
versity increased from the Upper Eocene to the Egerian
and decreased from the Egerian to the Ottnangian. In the
Karpatian, diversity slightly increased, followed by an
abrupt increase at the Karpatian/Badenian boundary. Fi-
nally, diversity decreased from the Moravian to the Sarma-
tian with an abrupt drop between the Badenian and the
From the FOs, appearance percentages and numbers of
FOs/1 Myr (Fig. 2B,C,D), the following intervals can be
characterized with high numbers of appearances of new
taxa: (1) The Karpatian and the Early Badenian (Moravi-
an) interval has most FOs at the Karpatian/Badenian
boundary, the Moravian has a high appearance percentage
while the Miocene peak of FOs/1 Myr falls in the Karpa-
tian; (2) The interval from Late Eocene to the Kiscellian/
Egerian boundary is characterized by a succession of less
significant appearance events, which led to the highest
number of benthic foraminiferal species in the Egerian.
The following boundaries are characterized by frequent
FOs: the Eocene/Oligocene, early/late Kiscellian and Kis-
cellian/Egerian, where a high appearance percentage is
to be found in the early Kiscellian and a high number of
FOs/1 Myr in the Late Eocene and early Kiscellian.
Appearance percentage reached the highest value in the
Sarmatian. This maximum does not correspond to a high
number of FOs; the high relative value is due to the low
number of species in the Sarmatian.
The intervals with high numbers of disappearing taxa
were distinguished using the LOs, extinction percentages
and number of LOs/1 Myr (Fig. 2E,F,G). (1) The interval
Late Badenian (Kosovian) and Sarmatian: the Kosovian
maximum of extinction percentage corresponds to the
highest number of LOs at the Badenian/Sarmatian bound-
ary. This maximum is preceded by the relatively high
numbers of LOs at the Moravian/Wielickian and the
Wielickian/Kosovian boundaries. The number of extinc-
tions/1 Myr reached the highest values in the Sarmatian;
(2) Concerning the Egerian: high values of LOs in the
Egerian and at the Egerian/Eggenburgian boundary, a
high extinction percentage as well as a high number of
LOs/1 Myr in the Egerian reflect a high number of disap-
peared foraminiferal species in the Egerian. (3) The num-
ber of extinctions/1 Myr reached the highest value in the
3.1.2 Planktonic species (Fig. 3)
Two maximum peaks from the Kiscellian to the Egerian
and in the Moravian characterize the species diversity
trend of planktonic species in the Central Paratethys
(Fig. 3A). As with the benthic Foraminifera, the diversity
of the Oligocene planktonic assemblages is higher than
those from the Miocene. In the Oligocene, diversity
reached a maximum value of 40 species in the Egerian,
while the highest Miocene diversity reached 30 species in
Three intervals with frequent first occurrence events
were distinguished (Fig. 3B,C,D): (1) The interval from the
Fig. 2. Species diversity trends of benthic foraminiferal species in the Central Paratethys, their FOs and LOs, appearance and extinction
percentages and number of FOs and LOs /1 Myr.
Late Eocene to the early Kiscellian is characterized by
high values of FOs in the Eocene and at the Eocene/Oli-
gocene boundary which corresponds to the high values of
appearance percentages and FOs/1 Myr in the Late Eocene
and early Kiscellian; (2) The interval from the Ottnangian/
Karpatian boundary to the Moravian is characterized by
high values of FOs at the Ottnangian/Karpatian and Karpa-
tian/Badenian boundaries and also with high values of ap-
pearance percentages in the Karpatian and Early Badenian
(Moravian). The highest value of FOs/1 Myr was recorded
in the Karpatian. (3) A minor peak of FOs, appearance per-
centage and FOs/1 Myr was recorded in the Egerian.
The highest number of disappearing species are concen-
trated in the following three intervals (Fig. 3E,F,G):
(1) The Late Badenian (Kosovian) and Sarmatian with
the highest values of LOs at the Badenian/Sarmatian
boundary, the highest extinction percentages in the Late
Badenian (Kosovian) and Sarmatian; (2) The Egerian with
a high value of LOs and extinction percentages; (3) A rela-
tively high value of LOs was also recorded at the early/
late Kiscellian boundary.
Species diversity patterns of benthic to planktonic fora-
minifers were compared from the Early Eocene to Oli-
gocene. The planktonic species appeared in two episodes:
from the Late Eocene to the early Kiscellian and in the
Egerian while new species of benthic foraminifers ap-
peared gradually in the whole interval. A higher number
of planktonic species disappeared at the early/late Kiscel-
For the Miocene, an increase in new planktonic species
started already at the Ottnangian/Karpatian boundary,
while the number of benthic species started to increase
only in the Karpatian.
3.1.3 Specific benthic groups (Fig. 4)
188.8.131.52 Agglutinated foraminifers (Fig. 4A,B,C,D)
The trend of species diversity changes of agglutinated
foraminifers (Fig. 4A) is similar to all benthic Foraminifera
except for the continuation of the Miocene species diver-
sity peak from the Moravian to the Wielickian. The high-
est species diversity of agglutinated foraminifers in the
FORAMINIFERAL SPECIES DIVERSITY IN THE CENTRAL PARATETHYS
Fig. 3. Species diversity of planktonic foraminiferal species in the Central Paratethys, their FOs and LOs, appearance and extinction
percentages and number of FOs and LOs/1 Myr.
Wielickian corresponds to the definition of the
Spiroplectammina carinata Zone by Grill (1941) in the
Middle Badenian. The trend also differs in a higher in-
crease in agglutinated foraminifers at the Ottnangian/Kar-
patian boundary and its rapid decrease between the
Wielickian and the Kosovian which agrees with the high-
est values of LOs at the Wielickian/Kosovian boundary
(Fig. 4D). The highest values of FOs are concentrated at
the early/late Kiscellian and the Karpatian/Badenian
boundaries (Fig. 4C).
The relative abundances of agglutinated foraminifers
(Fig. 4B) are comparable in the whole studied interval
(15—20 %) with exception of the Kosovian and Sarmatian
where relative abundance decreased under 10 % and the
Wielickian where agglutinated foraminifers reached the
maximum relative abundance (above 20 %).
184.108.40.206 Porcellanaceous foraminifers (Fig. 4E,F,G,H)
The species diversity of porcellanaceous foraminifers
concerning trends (Fig. 4E) are completely different from
the trends for all the benthic foraminifers. The number of
species is low (3—11) from the Late Eocene to the Karpa-
tian. At the Karpatian/Badenian boundary, the number of
species increases markedly (from 11 to 40 species) reach-
ing a plateau lasting until the end of Sarmatian. This coin-
cides with the highest value for FOs at the Karpatian/Bad-
enian boundary (Fig. 3G) and high values for FOs at the
Wielickian/Kosovian and the Badenian/Sarmatian bound-
aries. The highest value for LOs (Fig. 3H) was recorded at
the Badenian/Sarmatian boundary.
Relative abundance of porcellanaceous foraminifers
(Fig. 3F) increased from the Late Eocene to the Sarmatian:
they represented 5 % of benthic foraminiferal species in
the Eocene and Oligocene, 5—10 % in the Lower Miocene,
15—25 % in the Badenian and nearly 50 % in the Sarma-
220.127.116.11 Lagenids (Fig. 3I,J,K,L)
The lagenid species diversity has two peaks in the Ege-
rian and the Moravian (Fig. 3I). These correspond well to
the general species diversity pattern of all benthic fora-
minifers. Only ten species survived into the Sarmatian.
The highest number of FOs was recorded at the Karpatian/
Badenian boundary as for all benthic species, frequent
FOs were observed at the Eocene/Oligocene and the Kis-
cellian/Egerian boundaries (Fig. 4K). The most LOs are
to be found at the Badenian substage boundaries and at
the Egerian/Eggenburgian and the Badenian/Sarmatian
boundaries (Fig. 4L).
The relative abundances of lagenids (Fig. 4J) varied
around 20 % from the Late Eocene to the Moravian. These
values started to decrease only from the Middle Badenian
3.1.4 Groups with specific paleoecological require-
ments (Fig. 5)
The diversity trends of the groups with specific paleo-
ecological requirements (dysoxic, oxyphilic, euryhaline)
may reflect general paleoecological characteristics of the
18.104.22.168 Dysoxic species (Fig. 5A,B,C,D)
A bimodal distribution characterizes the species diversi-
ty values of the dysoxic species (Fig. 5A): the highest val-
ues are concentrated in the Kiscellian—Egerian interval.
The Karpatian-Badenian maximum is not so pronounced.
The highest number of FOs (Fig. 5C) was recorded at the
Eocene/Oligocene boundary, and there were high numbers
of LOs (Fig. 5D) at the Badenian/Sarmatian boundary and
in the interval from the Kiscellian/Egerian to the Egerian/
The relative abundances of dysoxic species (Fig. 5B)
vary around 20 % from the Kiscellian to the Badenian,
values above 20 % occur in the Oligocene, Karpatian and
in the Kosovian. The lowest value (about 5 %) was record-
ed in the Sarmatian.
22.214.171.124 Oxyphilic species (Fig. 5E,F,G,H)
The numbers of oxyphilic species (Fig. 5E) are low
(vary about 20—30) from the Eocene to the Karpatian/Bad-
enian boundary. Than the number markedly increased and
high values characterize the Badenian. This increase is ac-
companied by a high value of FOs (Fig. 5G) at the Karpa-
tian/Badenian boundary. After the highest number of LOs
at the Badenian/Sarmatian boundary (Fig. 5H), a slight de-
crease of oxyphilic species was recorded in the Sarmatian.
The relative abundances (Fig. 5F) vary from 10 to 20 %
from the Eocene to the Wielickian, while in the Sarmatian
it reached nearly 50 %.
126.96.36.199 Euryhaline (Fig. 5I,J,K,L)
The trend of species diversity changes for euryhaline
species (Fig. 5I) resembles that for porcellanaceous fora-
minifers and oxyphilic species with low values from the
Eocene to the Karpatian. The marked increase in number
of euryhaline species between the Karpatian and Badenian
is caused by the high number of FOs at the Karpatian/Bad-
enian boundary (Fig. 5K). Relative abundances of euryha-
line species (Fig. 5J) increased from 3 % in the Eocene to
about 70 % in the Sarmatian.
3.2 Species diversity events
The following foraminiferal species diversity events
were defined for the interval from the Late Eocene to the
Sarmatian in the Central Paratethys:
(1) The Egerian species diversity maximum (Figs. 2A,
3A, 4A,I and 5A). The species diversity of benthic species
reached a maximum due to constant increase in the num-
bers of foraminiferal species from the Eocene to the Egeri-
an (during approximately 15 Myr). This increase is the re-
sult of high numbers of FOs at the stage boundaries (the
highest was recorded at the Priabonian/Kiscellian bound-
ary) accompanied by no or low LOs from the Eocene to
the end of Kiscellian (Fig. 3B,E). The highest number of
lagenid species and a high number of textulariid species
were recorded in the Egerian. Dysoxic species reached the
maximum in the Egerian (Fig. 5A) but their species diver-
sity increased already from the Eocene/Oligocene bound-
The most planktonic foraminiferal species are found
during this Egerian maximum in the whole Central Para-
tethys, but their high number was already established in
the early Kiscellian (Fig. 3). The planktonic foraminiferal
species diversity reached the Egerian peak in two steps:
the first step represents the high numbers of FOs in the
Eocene and at the Eocene/Oligocene boundary. Numbers
of planktonic foraminiferal species were not lowered by
LOs in the interval from the Late Eocene to the early/late
Kiscellian boundary. The second step was the high num-
ber of FOs in the Egerian (20 % of new species appeared).
(2) The Egerian extinction. The increased foraminiferal
diversity in the Eocene and the Oligocene was interrupted
by a high number of LOs in the Egerian and at the Egeri-
an/Eggenburgian boundary (Figs. 2E, 3E, 4D, 5L,D). The
high total extinction corresponds to extinctions of the fol-
lowing groups: agglutinated foraminifers (39 %), lagenids
(33 %, especially at the Egerian/Eggenburgian boundary)
and dysoxic species (46 %). Also 33 % of planktonic spe-
cies became extinct.
(3) The Ottnangian species diversity minimum
(Figs. 2A, 3A, 4A,I, 5A) is characterized by a low number
of both planktonic and benthic species. In the terminal
Ottnangian Oncophora Beds only 25 indigenous benthic
species were recorded (Holcová 2001a). The low number
of species corresponds to low numbers of FOs of benthic
as well as planktonic foraminifers from the Eggenburgian
to the Ottnangian (i.e. during 3 Myr; Figs. 2B, 3B).
(4) The Early Badenian species diversity maximum
(Figs. 2A, 3A, 4I, 5I). The second peak of the bimodal spe-
cies diversity distribution curve does not reach the value
of the Egerian species diversity. The peak can be observed
for benthic and planktonic foraminifers as well as la-
genids. The agglutinated foraminifers as well as dysoxic
species (Figs. 4A, 5A) reached their second species diver-
sity peaks in the Wielickian, but the Moravian numbers of
species are very near to the Wielickian maxima.
The Early Badenian diversity peak is a result of an
abrupt increase in the number of benthic species at the
Karpatian/Badenian boundary (Fig. 2B): FOs became most
FORAMINIFERAL SPECIES DIVERSITY IN THE CENTRAL PARATETHYS
Fig. 4. Species diversity of some taxonomic groups (agglutinated foraminifers , porcellanaceous foraminifers, lagenids). Number of
species, relative abundances of each group and numbers of FOs and LOs are given.
Fig. 5. Species diversity of groups with specific paleoecological requirements (dysoxic, oxyphilic and euryhaline species). Number of
species, relative abundances of each group and numbers of FOs and LOs are given.
FORAMINIFERAL SPECIES DIVERSITY IN THE CENTRAL PARATETHYS
frequent in the history of the Central Paratethys at this
boundary. Numbers of lagenids and porcellanaceous fora-
minifers (Fig. 4G,K) as well as the numbers of oxyphilic
and euryhaline species (Fig. 5G,K) markedly increased at
this boundary. This increase of foraminiferal species diver-
sity agrees with the Early Badenian-build-up-event
(EBBE) recognized by Harzhauser & Piller (2007).
The species diversity of planktonic species increased
gradually from the Ottnangian/Karpatian boundary
(Fig. 3B); high numbers of FOs (and related appearance per-
centages) were recorded in the interval from the Ottnangian/
Karpatian boundary to the Early Badenian. During the Kar-
patian, the high values of appearance/1 Myr were recorded
for both benthic and planktonic species (Figs. 2D, 3D).
(5) Extinction at the Badenian/Sarmatian boundary is
shown by a severe drop in the number of foraminiferal spe-
cies. This extinction event was also defined by Harzhauser
& Piller (2007). The highest number of LOs were recorded
for benthic and planktonic foraminifers (Figs. 2E, 3E), por-
cellanaceous foraminifers (Fig. 4H), dysoxic, oxyphilic
and euryhaline species (Fig. 5D,H,L). Agglutinated fora-
minifers and lagenids became extinct in two steps. The
number of agglutinated foraminiferal species firstly de-
creased at the Wielickian/Kosovian boundary (Fig. 4C),
while the number of lagenids declined at the Moravian/
Wielickian boundary (Fig. 3L).
(6) The Sarmatian species diversity minimum (Figs. 2A,
3A, 4A,I, 5A,E). The lowest number of foraminiferal spe-
cies in the whole Central Paratethys history is in the Sar-
matian, as a consequence of the extinction events at the
Badenian/Sarmatian boundary. The number of Sarmatian
species represents only 29 % of the Egerian species and
49 % of the Ottnangian species. Extinction affected all
groups of benthic foraminifers. The marked decrease of
species diversity of agglutinated foraminifers and lagenids
was compensated by the appearance of new species of por-
Foraminiferal species diversity in a chain of periodical-
ly isolated and connected basins (as the Central Paratethys
was) was influenced by (1) foraminiferal species diversity
in the world’s oceans; (2) the diversity of biotopes settled
by foraminifers (presence/absence of different depth
zones, diversification of the environment in marginal fa-
cies); (3) the number of new immigrants and/or endemic
species dependant on paleogeographical events (isolation
vs. connection of basins); (4) the number of extinction
and/or disappearance events.
4.1 Paleogeographical events vs. species diversity
The following paleogeographical events (Fig. 6) were
compared with the foraminiferal species diversity changes
in the Central Paratethys:
(1) From the well communicating seas of the Eocene to
the early Kiscellian closures. The strait between the Medi-
terranean and the Paratethys was narrowed, and the Turgai
Strait, the Danish-Polish Trough and the Indic-Eastern
Paratethys connections were closed (Rögl 1998, 1999).
The first endemic Paratethys species appeared (Báldi
1986). Surprisingly, an increase of foraminiferal species
diversity was recorded during this first isolation event of
the Paratethys. This increase was represented partly by en-
demic species, and partly by new immigrants which ap-
peared in the surrounding marine basins and penetrated
through the narrowed gateways.
(2) After the end of this first isolation of the Paratethys,
a paleogeographically stable era followed of approximate-
ly 11 Myr (from the late Kiscellian to the Egerian/Eggen-
burgian boundary). During this period, the highest species
diversity was recorded with numbers of FOs relatively
high caused by penetration of new immigrants through the
new broad connections between the Central Paratethys
and the Mediterranean and the Indian Ocean.
To explain the cause of the end-Egerian extinction
event in a paleogeographically uniform period is difficult.
Oxygen concentration, salinity and sedimentological
characteristics changed only locally (Báldi & Seneš 1975;
Báldi 1986; Holcová 2001) and this local short-term oscil-
lation cannot have caused the Egerian extinction. Cooling
inferred from terrestrial biotopes by Planderová (1990)
agrees with the global Mi1 event (23.8 Ma) and can be ap-
proximately correlated with this extinction event. The in-
fluence of cooling in the Paratethys water masses was la-
tent because warm water masses penetrated from the
Indian Ocean into the Central Paratethys in the Egerian
(Rögl 1998, 1999). The probable cause of extinction may
be a decrease in nutrient availability. An indication of
these changes may be the acme of small-sized reticu-
lofenestras which characterizes the Oligocene/Miocene
boundary interval in the Central Paratethys (Holcová
2005). Small-sized reticulofenestras are considered to be
an indicator of warm water without high nutrient concen-
trations (Beaufort & Aubry 1992; Ćorić & Rögl 2004).
(3) The transgressive character of the Eggenburgian sed-
iments in many Central Paratethys basins (Steininger &
Seneš 1971; Kováč 2000) represents the base of the local
sea-level cycle CPC 1 which can be correlated with the
global cycle TB2.1 of Haq et al. (1988) (Kováč et al.
2001). This paleogeographical event is connected with
only a slight increase in the number of benthic species.
(4) Another important isolation of the Central Para-
tethys occurred during the Ottnangian (Papp et al. 1973).
The seaway between the Mediterranean and the Indian
Ocean was closed, and regressive, brackish facies ap-
peared. The decrease of foraminiferal diversity corre-
sponds to low values for FOs. In the terminal Ottnangian
Oncophora Beds, only 25 indigenous benthic foraminifer-
al species were recorded (Holcová 2001a).
(5) The Karpatian transgression terminating the Para-
tethys isolation (Brzobohatý et al. 2003) was accompanied
by a tectonic turnover and changes in the configuration of
the Central Paratethys basins. The transgression brought
an immigration of a new mollusc fauna (Rögl & Steininger
1983) and also an increase of the species diversity of
Fig. 6. Correlation between global paleoceanographical and foraminiferal events with Central Paratethys paleogeographical, paleoenvi-
ronmental and foraminiferal species diversity events.
FORAMINIFERAL SPECIES DIVERSITY IN THE CENTRAL PARATETHYS
planktonic foraminiferal species. The diversity of benthic
foraminiferal species only slightly increased.
(6) The Early Badenian large transgression established
normal marine facies in different depth zones (sublittoral
to bathyal) in the entire circum-Mediterranean area (Rögl
1998). The species diversity of benthic foraminifers in-
creased abruptly, probably caused by their quick immigra-
tion at the beginning of the large transgression.
(7) The Late Badenian marine flooding of the Central
Paratethys area (Rögl 1998) was connected with a persist-
ing broad communication with the Indo-Pacific. With the
persistence of this gateway, low-oxygen bottom condi-
tions were established, and the species diversity of fora-
miniferal assemblages decreased.
(8) The Sarmatian isolation was connected with the es-
tablishment of a very uniform paleoenvironment from the
Vienna to the Caspian Basins (Rögl 1998). The conven-
tional brackish water interpretation (e.g. Papp et al. 1974;
Rögl & Steininger 1983; Rögl 1998) was rejected by Pill-
er & Harzhauser (2005) but the large and abrupt faunal
turnover in the foraminiferal assemblages at the Badenian/
Sarmatian boundary indicates a quick and significant
change in the paleoenvironment between these two stages.
4.2 Foraminiferal species diversity events vs. paleoenvi-
ronment of the Central Paratethys
4.2.1 Sea-level changes
Generally, the conclusion of Schopf (1979) about the
correlation between sea-level oscillation and diversity was
also confirmed in the Central Paratethys. A global high-
stand period is replaced by a high species diversity. The
3rd-order sea-level cycle can be correlated with changes of
species diversity. The Egerian and the Lower Badenian
represented periods with the highest local Central Para-
tethys sea-level rise (Kováč & Hudáčková 1997; Kováč &
Zlinská 1998; Kováč et al. 1999, 2001) and the highest
species diversity. Sea-level fall (Ottnangian, Kováč &
Hudáčková 1997; Kováč & Zlinská 1998; Kováč et al.
1999, 2001) is related to lower species diversity.
The relation between regional Central Paratethys sea-
level changes and the global changes (Haq et al. 1988) has
been widely discussed (Rögl 1998; Kováč et al. 2001,
2004). The Ottnangian sea-level drop can be correlated
with a sea-level fall in the global cycle TB2.1, and the Sar-
matian one probably with TB2.6. The Early Badenian
transgression can be correlated with cycle TB2.3.
4.2.2 Climatic changes
The climatic changes in the Central Paratethys realm
and surrounding areas were studied mainly in the terrestri-
al biotops (Planderová 1990; Böhme 2003; Doláková &
Slamková 2003; Roth-Nebelsick et al. 2004; Kvaček et al.
2006). Paleotemperature changes in the marine basins
have been estimated from the geochemical data (Geary et
al. 1989; Šutovská & Kantor 1991; Krhovský et al. 1993;
Matyas et al. 1996; Durakiewicz et al. 1997; Hladilová et
al. 1998; Gonera et al. 2000; Bicchi et al. 2003; Latal et
al. 2004, 2006; Báldi 2006), from the composition of ma-
rine plankton (e.g. Spezzaferri & Ćorić 2001) and from the
presence of large foraminifers (Váňová 1975; Rögl &
Brandstätter 1993), development of algal-coral patch-reefs
(Pisera 1996), introduction of tropical mollusc faunas and
fish (Steininger et al. 1978; Bellwood & Schultz 1991).
The Late Eocene to Early Oligocene cooling event was
one of the causes of the Early Oligocene thermohaline water
stratification (Popov et al. 2004) followed by the Late Oli-
gocene warming event (Roth-Nebelsick et al. 2004). These
two events did not affect the gradual increase in the number
of foraminiferal species from the Early Eocene to the Egeri-
an leading to the Egerian species diversity maximum.
The influence of the cooling event at the Oligocene/Mi-
ocene boundary (which can be correlated with the global
Mi1 event, Billups & Schrag 2002) on the temperature of
the Paratethys sea was perhaps latent (see Chapter 4.1).
The Eggenburgian warming event and the Late Ottnan-
gian-Early Karpatian cooling event (Planderová 1990;
Doláková & Slamková 2003) did not substantially influ-
ence foraminiferal species diversity.
The Middle Miocene Climatic Optimum and the follow-
ing cooling event (14—13.5 Ma; comparable with Mi3
event), have also been studied in detail in the Central
Paratethys (Schwarz 1997; Gonera et al. 2000; Böhme
2003; Bicchi et al. 2003; Báldi 2006). Temperature
changes can be associated with marine circulation inde-
pendently in both the oceanic realm and in the Central
Paratethys (Báldi 2006). In the Central Paratethys, approx-
imately after the Middle Miocene, a Climatic Optimum
was probably connected with a salinity crisis. The fora-
miniferal assemblages reflected these environmental
changes by maintenance of overall high species diversity
and the highest diversity of agglutinated foraminifers in
the Middle Badenian. A change of antiestuarine to estua-
rine circulation took place at the Middle/Late Badenian
boundary (Báldi 2006). The change of circulation led to
stratification of the water column in the Late Badenian. In
some deep parts of the Central Paratethys basins stagna-
tion and accumulation of dysoxic and anoxic sediments
occurred (Báldi 2006). Benthic foraminiferal diversity de-
creased in the Late Badenian.
4.2.3 Oxygen content
Two dysoxic intervals with uniform disaerobic bottom
conditions in the entire Central Paratethys were described
in the early Kiscellian and the Late Badenian (Papp et al.
1978; Báldi 1986; Kováč 2000; Smith et al. 2001).
Dysoxic facies were recorded locally in the late Kiscel-
lian, Egerian, Eggenburgian, Ottnangian, Karpatian, Mid-
dle Badenian and early Sarmatian (Steininger & Seneš
1971; Papp et al. 1973; Báldi & Seneš 1975; Papp et al.
1978; Steininger et al. 1985; Kováč et al. 1989; Holcová-
Šutovská 1996; Kováč 2000; Holcová 2001b; Brzobohatý
et al. 2003). With the exception of the Ottnangian (Papp et
al. 1973; Kováč et al. 1989), dysoxic environments were
inhabitated by benthic foraminifers.
Oxic bottom condition with no or minimal dysoxic fa-
cies characterized the Early Badenian and the late Sarma-
tian (Papp et al. 1974, 1978).
The following changes in foraminiferal assemblages can
be correlated with changes in bottom oxygenation:
(1) A high number of FOs corresponding to a marked in-
crease in the number of dysoxic benthic species (Bolivina
spp., non-costate Bulimina spp., Uvigerina spp., Fursen-
koina spp.) was recorded between the Eocene and Oli-
gocene together with the establishment of uniform disaer-
obic bottom conditions.
(2) Dysoxic intervals (the early Kiscellian and the Late
Badenian) are connected with lower numbers of aggluti-
In the Sarmatian dysoxic species were very rare and the
relative abundance of oxyphilic species was high.
4.2.4 Salinity oscillations
A generally accepted hyposaline condition is found in
the history of the Central Paratethys only in the late Ott-
nangian (Papp et al. 1973; Rögl 1998). Decrease of salini-
ty was connected with a minimum number of foraminiferal
taxa. The number of euryhaline species stayed constant.
The diversity of euryhaline species increased markedly
at the Karpatian/Badenian boundary but this event is not
connected to a bottom salinity reduction. An abrupt in-
crease in the relative abundance of euryhaline taxa con-
nected with a faunal turnover at the Badenian/Sarmatian
boundary has been conventionally explained by a hy-
posaline event. This interpretation was rejected by Latal et
al. (2004) and Piller & Hauzhauser (2005). However,
marked changes in foraminiferal assemblages indicate pa-
leoenvironmental changes connected with the isolation of
Central Paratethys foraminiferal species diversity events
vs. global events
(1) The Late Eocene foraminiferal extinction (Kaiho
1994) occurring between 38 and 34 Ma (Boersma 1986;
Keller 1986; Keller et al. 1992) was isochronous with the
“birth” of the Paratethys (Rögl 1998, 1999; Popov et al.
2004) and did not influence the species diversity of fora-
minifers in the Central Paratethys. The Oligocene increase
of foraminiferal diversity in the Central Paratethys may
have been influenced by the Late Oligocene post-extinc-
tion foraminiferal speciation in the surrounding basins
(e.g. Kaiho et al. 1993; Hernitz Kučenjak et al. 2006). Pen-
etration of new immigrants was possible through the new
broad connections between the Central Paratethys and the
Mediterranean and the Indian Ocean (see Chapter 4.1).
(2) A moderate Late Oligocene (27—25 Ma) low-oxygen
event connected with the extinction of deep-water fauna
was documented by Kaiho (1994) in the world ocean. This
can be correlated with the early Egerian but no extinction
event was observed at this time in the Central Paratethys.
(3) The early Middle Miocene extinction interval
(Kaiho 1994) was dated between 17—13 Ma (Thomas
1986; Thomas & Vincent 1987; Berggren & Miller 1989)
and can be correlated approximately with the Karpatian
and Badenian (Cicha et al. 1998; Hohenegger et al. 2007).
The causes of the event have been explained by water
mass changes resulting from warming (associated with
sluggish circulation) and subsequent cooling in high-lati-
tude surface and deep water (Thomas & Vincent 1987;
Woodruff & Savin 1991). These climatic changes (Climat-
ic Optimum and subsequent cooling) also influenced the
local circulation in the Central Paratethys sea and led to
gradual benthic foramininiferal extinction in the Middle
and Late Badenian.
Six foraminiferal species diversity events can be defined
in the Central Paratethys:
1 – The Egerian species diversity maximum was
reached by the progressive appearance of new foraminifer-
al species from the Eocene to the Egerian (approximately
15 Myr). Up to the Egerian this increase of species diversi-
ty was accompanied by no or minimal disappearances of
foraminiferal species. The early Kiscellian isolation of the
Central Paratethys did not interrupt this increase of spe-
2 – The large Egerian extinction is difficult to explain
because the Egerian represented a long interval with gen-
erally uniform paleoecological and paleogeographical
conditions. The extinction is isochronous with Mi1 cool-
ing event. The acme of small-sized Reticulofenestra repre-
sents a regional isochronous event which may indicate a
decrease of nutrient availability at the Oligocene/Miocene
3 – The Ottnangian species diversity minimum
evolved gradually from the Eggenburgian (approximately
3 Myr) by constant disappearance of foraminiferal species
without compensation from the appearance of new spe-
cies. This was a consequence of the Paratethyan isolation,
deterioration of marine environments and development of
anoxic and hyposaline biotopes without foraminifers.
4 – The Moravian species diversity maximum was con-
nected with an abrupt input of benthic immigrants during
a transgression. Planktonic foraminiferal species diversity
started to decrease from the Ottnangian/Karpatian as a
consequence of the opening of the new sea-ways.
Influence of the global cooling event Mi3 on the Para-
tethyan circulation in the Late Badenian is supposed. This
circulation change influenced a gradual decrease of fora-
miniferal species diversity.
5 – The extinction and faunal turnover at the Bade-
nian/Sarmatian boundary indicates quick and significant
change of paleoenvironments between these two stages.
6 – The Sarmatian species diversity minimum shifted
the composition of assemblages to the endemic euryhaline
species. These changes can be clearly correlated with the
isolation and vanishing of the Central Paratethys basins.
Three of the above mentioned foraminiferal species di-
versity events (the Ottnangian minimum, the Moravian
FORAMINIFERAL SPECIES DIVERSITY IN THE CENTRAL PARATETHYS
maximum and the Sarmatian minimum) were influenced
mainly by the regional isolation/communication paleo-
geographical events. However, the Ottnangian and Sarma-
tian isolation as well as the Early Badenian transgression
may be related to global sea-level changes. The decrease
of species diversity in the Late Badenian can be explained
by regional paleoenvironmental changes provoked by
global cooling. The Egerian extinction cannot be satisfac-
torily explained, at this moment.
Acknowledgment: The author wishes to thank to review-
ers (S. Spezzaferri from Fribourg University, W.E. Piller
from the University of Graz, K. Báldi from Budapest Uni-
versity) for their comments, which greatly improved the
manuscript during the review process. The study was sup-
ported by the Project MSM0021620855.
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