GeolCarp_Vol60_No3_251

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA, JUNE 2009, 60, 3, 251—262 doi: 10.2478/v10096-009-0017-0

Introduction

The present study shows the results of ostracod analyses
from the Sadovi section (Mt Požeška gora, Croatia).

During the Early Miocene this area was located in the cen-

tral  part  of  the  North  Croatian  Basin,  at  the  south-western
margin of the Pannonian Basin System (Fig. 1, Fig. 1.1). The
formation of the North Croatian Basin (in further text NCB)
was  connected  with  passive  continental  rifting.  A  syn-rift
phase began during the Ottnangian and lasted until the Mid-
dle  Badenian.  The  post-rift  phase  lasted  from  the  Middle
Badenian to the Pliocene (Pavelić 2001).

In the NCB brackish ostracod assemblages are common in

Upper Miocene deposits (Sokač 1961, 1961a, 1963, 1965,
1967, 1972), but the presence of brackish ostracod faunas in
Lower Miocene deposits (Upper Ottnangian/Lower Karpa-
tian) at Sadovi locality is a new discovery.

The appearance of such fauna indicates the first ingression

of marine water into the Late Ottnangian/Early Karpatian
lake. In the uppermost part of the section Karpatian marine
marls concordantly overlay these brackish marls.

Geological setting

The Lower Miocene (Upper Ottnangian/Lower Karpatian)

deposits of the investigated section belong geotectonically to
the Pannonian Basin System and paleogeographically to the
south-western  margins  of  the  Central  Paratethys.  Central
Paratethys extends from Bavaria to the Carpathian mountain
chain  (Steininger  &  Rögl  1979,  1984;  Rögl  &  Steininger
1983, 1984; Rögl 1998, 1999). It was a part of the interconti-
nental  bioprovince  which  began  to  evolve  in  the  Oligocene
and was formed due to collision of the European (Tisa-Moe-
sia) and the African Plates (Horváth & Royden 1981; Kováč
et al. 1998).

Early Miocene ostracods from the Sadovi section

(Mt Požeška gora, Croatia)

VALENTINA HAJEK-TADESSE

, MIRKO BELAK

, JASENKA SREMAC

, DAVOR VRSALJKO

and LARA WACHA

Croatian Geological Survey, Sachsova 2, 10 000 Zagreb, Croatia;

valentina.tadesse@hgi-cgs.hr

Faculty of Science, Department of Geology, Institute of Geology and Paleontology, Horvatovac 102 a, 10 000 Zagreb, Croatia

(Manuscript received November 8, 2007; accepted in revised form October 23, 2008)

Abstract: The study of the Early Miocene (Late Ottnangian/Early Karpatian) ostracod fauna from the Sadovi section
(Mt Požeška gora, Croatia) led to several results concerning Neogene paleobiogeography and paleoecology. Brackish
deposits of Late Ottnangian and Early Karpatian age have been recognized for the first time in the North Croatian Basin.
These deposits indicate the first marine ingression into the Early Miocene lake in this area. Twenty-nine ostracod species
were determined, including the new taxa Fabaeformiscandona slavonica nov. sp. and Herpetocypris sadovii nov. sp.

Key words: Late Ottnangian, Early Karpatian, North Croatian Basin, paleoecology, brackish environment, Ostracoda.

During the Miocene, connections of the Central Paratethys

with the Mediterranean and the Indo-Pacific Ocean were es-
tablished  and  interrupted  several  times  (Steininger  et  al.
1988; Rögl 1996). Such unstable type of connection resulted
in  sea-level  oscillations  (Haq  1991),  different  depositional
modes  and  paleoecological  conditions,  forcing  the  develop-
ment of different ostracod faunas.

In the Central Paratethys the Lower Miocene Ottnangian

stage (~ 18—17 Ma) is a twofold stage (Rögl et al. 1973). Ma-
rine conditions were dominant in the early phase of the Ott-
nangian. Except for the Northern Alpine Foreland Basin, no
entirely marine environments are known (Harzhauser & Piller
2005, 2007). During the Late Ottnangian, the Paratethys was
isolated  from  the  Mediterranean  Sea.  This  sea-level  drop
seems  to  be  linked  with  the  global  sea-level  change  of  the
TB2.1 cycle, lasting from ~ 21—17.5 Ma (Haq 1991). Brack-
ish to freshwater sedimentary conditions prevailed, resulting
in the deposition of Rzehakia/Oncophora Beds in large areas
of  the  Paratethys  (Rzehak  1882;  Rögl  &  Steininger  1983;
Rögl 1998).

At the beginning of the late Early Miocene, during the glo-

bal  sea-level  cycle  TB  2.2,  lasting  from  ~ 17.5—16.4 Ma
(Haq  1991),  the  sea-level  rose  and  the  Karpatian  stage
(~ 17—16.3 Ma)  began  with  transgressions.  In  different  ba-
sins,  Karpatian  deposits  were  discordantly  deposited  over
the  Ottnangian  marine  shales,  in  littoral  settings  over  the
Rzehakia  Beds, continental deposits or over older basement
units (Rögl et al. 2003). A continuous transition between the
Ottnangian  and  the  Karpatian  is  expected  only  in  deeper
parts  of  the  Central  Paratethys,  whereas  all  shallow  marine
settings display a strong discordance (Rögl et al. 2003).

In the NCB the Lower Miocene lowstand deposition is

represented  mainly  by  alluvial,  fluvial  and  deltaic  deposits.
At  the  base  of  the  Karpatian,  this  sequence  is  rapidly  re-
placed  by  marine  neritic  to  shallow  bathyal  sediments
(Pavelić 2001).

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HAJEK-TADESSE, BELAK, SREMAC, VRSALJKO and WACHA

Formation of a sedimentary basin in the area of the

Slavonian Mts started in the Ottnangian with freshwater de-
posits  (Pavelić  et  al.  1998).  Two  depositional  phases  were
recognized by Pavelić (2001). The first phase was character-
ized by the accumulation of predominantly breccias and con-
glomerates with subordinate sand and siltstones deposited in
a  braided  alluvial  plain  and  under  semi-arid  climate  condi-
tions  (Pavelić  et  al.  1998).  During  the  second  depositional
phase  a  freshwater  lake  was  formed  in  the  area  of  the
Slavonian Mts, in which silts and sands with sporadic layers
of gravels, tuffs and tuffites were deposited. The endemic os-
tracod  and  molluscs  faunas  are  found  together  with  fossil
plants  (Pavelić  et  al.  1998).  The  second  depositional  phase
was  characterized  by  a  shift  to  humid  climate  conditions.
Lacustrine conditions were replaced by marine environments
during the Karpatian time (Pavelić 2001).

Study site and methods

Study site

The Sadovi section is located in

the  southern  part  of  Mt  Požeška
gora  on  the  top  of  the  Sadovi
hills, along a forest road. The sec-
tion is about 500 m long, and the
total  real  thickness  of  the  Lower
Miocene  sequence  is  150 m.  All
32  samples  were  collected  from
marls  (Fig. 2).  Besides  the  ostra-
cods,  remains  of  plants,  molluscs
and fishes were found. The colour
of  the  marls  varies  from  yellow,
brownish-grey to olive-green.

In the first 30 meters of the sec-

tion,  the  marls  are  laminated  and
rich  in  plant  fragments,  ostracods
and  fish  bones.  In  the  central  part
of the section (65—80 m) the marls
are laminated, contain thin films of
bitumen  and  rich  ostracod  faunas.
Twenty  five  meters  of  the  section
are not exposed. Between samples
Sa 29  and  Sa 30,  the  contact  with
the  Karpatian  marine  sediments
has been noticed.

Methods

The collected marl samples were

disaggregated  by  soaking  in  a  hy-
drogen  peroxide  solution  for  24
hours, then washed through sieves
(0.5;  0.25;  0.125;  0.063 mm)  and
dried. Some of the samples needed
extra  cleaning  and  were  soaked
again  in  hydrogen  peroxide  and
treated  ultrasonically  for  approxi-
mately  20  seconds.  100 g  of  each
dried residue was observed under a

Fig. 1. Geographic setting of the North Croatian Basin within the Pannonian Basin System (after
Pavelić 2001). Fig. 1.1. Location map of Mt Požeška gora

with marked position (•) of the Sa-

dovi section (after Pavelić 2001).

stereomicroscope. Ostracods and other remains (foraminifers
and gastropods) were hand-picked, counted and determined.
Ostracods were picked qualitatively but not selectively, in or-
der to preserve the relative composition of the thanatocoenose.

Six samples were selected for insoluble residue and car-

bonate content analysis. They were treated with 10% HCl
and the carbonate content was measured. The insoluble resi-
due, after treatment with HCl, was washed and thin sections
were prepared.

Photographs were made with a SEM at the Geological De-

partment of the Faculty of Science in Zagreb. For photo-
graphing we used two different detector types: BSE – Back
Scatter Detector and SE – Secondary Electron Detector.

All ostracod specimens are stored in the archive collection

of the Croatian Geological Survey (Inv. No. HGI-CGS Hr/Sa
2006/HTV).

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EARLY MIOCENE OSTRACODS FROM THE SADOVI SECTION (CROATIA)

Results

Ostracod distribution

Altogether, 29 species belonging to 18 genera and 5 fami-

lies (Darwinulidae, Candonidae, Cyprididae, Limno-
cytheridae and Cytherideidae) were identified. Some of the
taxa (13) remain in open nomenclature, 6 species are left in
open nomenclature, two species are new (Table 1).

Fig. 2. Stratigraphical log of the Sadovi section.

A brackish ostracod fauna was found in 26 samples. Only

the uppermost three samples (Sa 30—32) of the section repre-
sent a marine fauna; we recognized juvenile valves of marine
Karpatian  ostracods  (Henryhowella  sp.),  and  benthic  and
planktonic  foraminifers  (Svratkina  sp.,  Coryphostoma  cf.
digitalis, Tenuitella sp., Globigerinoides sp., Cassigerinella
globulosa).  This  environmental  change  was  caused  by  the
influx of sea water into the lake. The marine depositions will
not be part of this investigation. Three samples (Sa 4, Sa 5,
Sa 14) did not yield any fossils.

The most abundant and frequent species in the samples

from the Sadovi section are: Herpetocypris sp. (22 records),
Candona (Pontoniella) sp. and Candona sp. (both 13 records)
and Fabaeformiscandona pokornyi (11 records). The diversity
varies throughout the section, and is generally low.

The maximum number of species (11) was reported in

samples Sa 2 and Sa 9. Other samples with relatively high
numbers of species are Sa 18 (10 species), Sa 7 (9 species)
and Sa 1, Sa 10, Sa 28 (8 species). In other samples the num-
ber of species varies between 4 and 8, and 5 samples contain
only 2 species.

The identified ostracod species (Table 1) can be divided

into two groups. The first group includes the species from
samples 1 to 18 of Sadovi section. These samples point to a
brackish environment with salinities ranging from oligoha-
line or (low) mesohaline.

The second group is represented by the species from sam-

ples 19 to 29, which indicate the gradual decrease of salinity.

A few species appear exclusively in the first group: Fa-

baeformiscandona  slavonica  nov.  sp.,  Cyprideis  sublittora-
lis, Heterocypris cf. francofurti, Amplocypris sp., Cypris sp.,
Darwinula  stevensoni,  Eucypris  cf.  dulcifons,  Paralimno-
cythere  rostrata,  and  three  Potamocypris  species:  P.  fulva,
P. cf. gracilis, P. cf. arcuata. The accompanying species are
Fabaeformiscandona    pokornyi,  Candona  (Caspiolla)  sp.,
Candona (Pontoniella) sp., Cypridopsis biplanata, Herpeto-
cypris  sp.  and  Herpetocypris  sadovii  nov.  sp.  In  the  upper-
most  part  of  the  first  level  Candona  (Lineocypris)  sp.  and
Candona  (Typhlocypris)  sp.,  have  the  first  common  occur-
rence.  A  single  valve  of  Aurila  sp.  and  a  few  benthic  fora-
minifers Aubignyna perlucida   were found in sample Sa 2.

The second group of samples (from Sa 19 to Sa 29) con-

tains most of the species from the first group. The dominant
species in the second group are Herpetocypris sp., Fabaefor-
miscandona pokornyi and Candona (Pontoniella) sp.

Herpetocypris sp. gradually becomes more common from the

end of the first level to the end of the section. The species which
are present only in the second group with very few valves are
Pseudocandona praecox, Mediocypris cf. candonaeformis and
Dolerocypris  sp.  In  contrast  to  the  first  group  Candona
(Typhlocypris)  sp.  is  more  common  in  the  second  group.
Candona  (Lineocypris)  sp.,  Candona  (Caspiolla)  sp.,  Cypria
sp. and Herpetocypris sadovii nov. sp. occur only in the upper-
most  samples  of  the  second  group.  Cypria  dorsalta,  Cypri-
dopsis biplanata and Eucypris sp. are present with few valves.

The brackish ostracod fauna is autochthonous, and well

preserved. The population structure of the ostracod fauna in-
dicates low water energy of the environments and low sedi-
mentation rates (Whatley 1983).

254

HAJEK-TADESSE, BELAK, SREMAC, VRSALJKO and WACHA

Mineralogical analyses and carbonate content

Carbonate content in marl samples SA 4, SA 7, SA 14,

SA 16 and SA 19 varies from 46.3 to 55.0 %. Sample SA 30
is  a  clay  rich  marl  with  13.1%  carbonate  content.  Samples
from Sadovi contain a very small amount of silty siliciclastic
detritus. The mineral composition of the insoluble residue in
the  analysed  samples  is  quite  homogeneous.  Non-undulose
monocrystalline subrounded or subangular quartz grains and
altered  feldspar  grains  dominate.  Non-altered  feldspars  and
twinned  plagioclase  are  less  abundant.  The  lithic  fragments
are  scarce,  with  domination  of  shale  fragments  and  some
chert fragments.

A very small amount of heavy minerals is present in the

samples. The most abundant among them are resistent tur-
maline and garnet, accompanied by rutile, zircon, staurolite,
amphibole, titanite, biotite and chlorite. Opaque minerals are
also present. In the sample SA 19 opaque minerals dominate,

while the composition of the translucent heavy minerals is
the same as in other samples.

The source of clasts could be related to igneous and meta-

morphic rocks from the vicinity. The small amount of silici-
clastic components indicates calm depositional conditions
and a very restricted terrigenous input.

Discussion

Ostracods from brackish waters are valuable tools in paleo-

environmental  reconstruction  and  in  environmental  studies.
The main parameters which can be monitored with brackish
water  ostracods  are  salinity  and  hydrochemistry,  water
depth,  substrate,  oxygen  and  productivity,  temperature  and
climate (Frenzel & Boomer 2005). The composition of ostra-
cod assemblages from our section depends mainly on the sa-
linity and depth of the lake water.

Table 1: Distribution of the Ostracoda in the Lower Miocene deposits of the Sadovi section (1—18 = transgressive phase; 19—29 = regres-

sive phase). Only adult valves for each species in the samples were counted:

1—2,

2—5,

5—10,

10—20,

20—40.

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EARLY MIOCENE OSTRACODS FROM THE SADOVI SECTION (CROATIA)

Brackish-water ostracods are normally found in transitional

environments  or  in  inland  saline  lakes  (Remane  1971).  The
different biotopes appear and disappear at various levels in a
stratigraphic succession, following the dynamics of transgres-
sion and regression through time (Do Carmo et al. 1999). In
the Western Paratethys, the Late Ottnangian regression initiat-
ed the development of a shallow brackish basin, and presence
of  planktonic  larval  development  in  gastropods  indicates  a
connection  with  the  open  sea  (Kowalke  &  Reichenbacher
2005). In the Sadovi section deposition of Upper Ottnangian
and  Lower  Karpatian  brackish  water  sediments  is  a  conse-
quence of a marine ingression into the freshwater lake, which
is the first signal of the new transgression in the Karpatian.

During the Early Ottnangian a freshwater lake existed in the

area of the Mt Požeška gora (Kochansky-Devidé 1979;
Pavelić et al. 1998; Pavelić 2001). The oldest alluvial and lake
clayey deposits lack ostracods, and marly sediments of a olig-
otrophic freshwater lake contain ostracod communities
(Hajek-Tadesse et al. 2006).

The increase of water salinity in the Late Ottnangian and

Early Karpatian, which is observed in the Sadovi section for
the first time, was caused by the ingression of marine water
into the lake. Ingression of marine water was documented by
the composition of ostracod fauna, especially with the occur-
rence of typical brackish ostracod taxa: Cyprideis sublittora-
lis and Aurila sp. together with scarce foraminifers
(Aubignyna perlucida).

The occurrence of the genus Cyprideis in the Sadovi sec-

tion plays an important role in the identification of brackish
environments.  The  genus  Cyprideis  is  not  typical  for  the
Lower Miocene deposits of the Central Paratethys. It is com-
mon in the Upper Miocene deposits of Croatia (Sokač 1972).
For the Late Sarmatian and Pannonian (12 to 8.5 Ma) in the
Molasse  Basin,  the  occurrence  of  Cyprideis  spp.  and  can-
donid  species  indicate  brackish  conditions  (Janz  &  Venne-
mann 2005).

In Europe the probable ancestor of the genus Cyprideis is

Miocyprideis (Kollmann 1960). The evolutionary sequence of
Cyprideis in the Amazon Basin began in the Early Miocene,
and  the  most  explosive  radiation  of  Cyprideis  in  Western
Amazon  region  occurred  at  the  boundary  between  the  Early
and Middle Miocene (Langhian stage). The ancestors of Cyp-
rideis in the Amazon Basin are not identified. A paleofauna of
21 endemic species of the euhaline genus Cyprideis lived be-
tween  polyhaline  and  oligohaline  salinity  ranges  during  spo-
radic intervals of marine influence (Mu  oz-Torres et al. 2006).

In contrast to brackish Rzehakia/Oncophora Beds of the

Central Paratethys, which is a result of global sea-level de-
crease, Lower Miocene brackish deposits in Sadovi are the re-
sult of marine ingression.

Numerous paleosalinity studies were based on the cosmopol-

itan holoeuryhaline species Cyprideis torosa. Aladin (1993)
pointed out that C. torosa is a brackish water ostracod of marine
origin with extremely complex osmoregulatory mechanisms
and is known to withstand low oxygen conditions.

Lateral sieve pores of recent and fossil Cyprideis torosa

vary in outline from round to oval and irregular. The percent-
age of round pores decreases with the increase of salinity
(Rosenfeld & Vesper 1977).

Gliozzi & Mazzini (1998) and Anadon et al. (2002) note

that differences in paleosalinity values may be explained by
mixing waters of different origin.

A connection between the salinity and noding of C. torosa

was noticed by Hartmann (1964), Vesper (1975), Van Harten
(1996) and Keyser (2005). Noding at low salinities is the most
accepted condition, despite the fact that the controlling mech-
anism is poorly understood. Do Carmo et al. (1999) connect
variable noding of Middle Jurassic limnocytherid ostracods
with salinity changes.

In the first group (transgressive part) of the Sadovi samples,

rounded, oblong and irregularly shaped sieve pores of the spe-
cies Cyprideis sublittoralis are distributed in almost even per-
centage.  Rosenfeld  &  Vesper  (1977)  connected  such
distribution  with  8  to  16 ‰  salt  concentration.  On  the  other
hand, in these samples we found only unnoded forms of adults
and  numerous  juvenile  stages.  Brackish  water  ostracods  live
within  the  uppermost  few  millimeters  of  the  sediment,  be-
cause much of the sediment below this level is often rich in or-
ganic material and, as a result, depleted in oxygen (Horne &
Boomer 2000).

The ostracod fauna from Sadovi can be compared with the

brackish  ostracods  from  the  Kirchberg  Formation  (Upper
Ottnangian/Lower  Karpatian)  of  the  western  part  of  the
South  German  Molasse  Basin  (Witt  2000)  and  ostracods
from the Oncophora Beds in the East (Witt 1999), but ostra-
cod  assemblages  from  Sadovi  are  indicative  for  a  more  sa-
line  lake environment.

Such faunal relations are more common in the Late Mi-

ocene of the Paratethys area: Styrian Basin, Turčianska kotli-
na Depression (Slovakia) and the Pannonian Basin (Sokač
1972; Gross 2004; Pipík & Bodergat 2004).

Beside salinity, substrate is the main factor controlling the

occurrence of ostracods in brackish water (Ruiz et al. 2000).
The marl deposits in the Sadovi section are horizontally lami-
nated, they contain thin bituminous lamina, rich ostracod as-
semblages, plant remains and a few skeletal fish particles.

During the Late Ottnangian and Early Karpatian in the area

of Mt Požeška gora an eutrophic lake existed with increasing
salinity of water and low input of clastic material from the
land. We recognized this event as the new, “third phase” of
deposition in NCB sensu Pavelić (2001).

Most of the ostracod taxa determined from the Sadovi sec-

tion prefer ponds and lakes and tolerate an increase of salinity
(Meisch 2000). Dissolution of valve margins in the species
Amplocypris sp. and bacterial activities on Cypria dorsalta
valves are a common effect and are typical for eutrophic envi-
ronments (Reeves et al. 2007).

Carbonel et al. (1988) conclude that species with “high” tri-

angular, trapezoidal shape and elongated valves with pointed
posterior (such as Candona (Lineocypris) sp., Candona
(Typhlocypris) sp., Candona (Pontoniella) sp. and Cypris sp.
from Sadovi) occur frequently in persistent environments with
low environmental fluctuations like groundwater habitats and
the sublittoral of old lakes.

The confirmation of marine ingression into the Late Ott-

nangian/Early Karpatian lake in the area of Mt Požeška
gora gives a key for understanding the formation of analci-
me deposits north and south-east from the investigated sec-

256

HAJEK-TADESSE, BELAK, SREMAC, VRSALJKO and WACHA

tion. Ščavničar et al. (1983) studied the analcime-bearing
deposits north of Sadovi in Poljanska (southern Papuk,
Slavonija region) and Sirječić et al. (1974) described analci-
molites south-east from Sadovi, in the Tuzla salt series (Bos-
nia and Herzegovina).

The genesis of sediments near Poljanska is linked to the

saline  alkaline  lake  environment.  Ščavničar  et  al.  (1983)
conclude that analcime was formed by alternation of volca-
nic glass, as well as from other silicates in the shallow water
of an unsheltered saline alkaline lake, under conditions of in-
creasing  aridity  and  closure  of  the  lake.  Variable  composi-
tion, texture and structure of analcime-bearing deposits give
the  evidence  of  the  climate  shifts  and  the  lake  history
(Ščavničar et al. 1983).

The registered influx of marine water into the lake before

the Karpatian transgression and the appearance of the brack-
ish ostracod fauna at Sadovi raise new questions on marine
connections between the Mediterranean and the Central
Paratethys.

The problem of the marine connection between the Medi-

terranean and Central Paratethys area remains unsolved. The
Transtethyan Trench Corridor (Bistričić & Jenko 1985) was
situated somewhere in the territory now forming part of Slo-
venia  or  northern  Croatia.  On  the  basis  of  our  previous  in-
vestigations  and  the  results  of  our  work  in  this  paper  we
conclude  that  the  possible  position  of  the  new  corridor,
which  has  not  previously  been  well  documented,  is  south-
east of the NCB.

Conclusions

The study of ostracod assemblages from Sadovi revealed

for  the  first  time  the  existence  of  brackish  deposits  in  the
Late  Ottnangian/Early  Karpatian,  as  a  result  of  marine  in-
gression into the lacustrine water of the NCB. In the upper-
most  samples  of  the  investigated  section  a  continuous
transition  of  brackish  deposits  into  the  Karpatian  marine
beds with scarce marine microfauna was observed.

The microfauna from the Sadovi section belong to the up-

per  part  of  the  Lower  Miocene  (Upper  Ottnangian/Lower
Karpatian).  A  more  precise  biostratigraphic  dating  is  not
possible on the basis of ostracods. However, paleoecological
data  obtained  by  ostracod  analysis  are  much  more  signifi-
cant.  For  the  first  time,  based  on  ostracod  assemblages,  a
wide brackish influence could be documented in the Lower
Miocene deposits of the NCB.

Samples from the Sadovi section can be divided into two

groups. The groups are defined on the basis of different os-
tracod assemblages and different trends in water salinity. In
the  first  group  (samples  1—15)  we  found  brackish  ostracod
assemblages  with  the  indicative  brackish  species  Cyprideis
sublittoralis, which has been detected for the first time in the
Lower  Miocene  of  the  Central  Paratethys.    In  the  second
group (samples 16—25) these species disappear and ostracod
assemblages indicate the slight decrease of water salinity.

Qualitative analyses resulted with 29 identified ostracod

species, among which two species are new: Fabaeformiscan-
dona slavonica nov. sp. and Herpetocypris sadovii nov. sp.

Brackish deposits of the Sadovi section are the result of

the marine ingression into the lake. They represent the “third
phase” of development, of the NBC successively comple-
menting the local reconstruction of the Ottnangian deposi-
tional model introduced by Pavelić (2001).

The genesis of analcime layers north and south-east of the

investigated area (Sirječić et al. 1974; Ščavničar et al. 1983)
can also be explained by the intrusion of sea water into the
Early Miocene lake.

The detected brackish Lower Miocene (Upper Ottnangian/

Lower  Karpatian)  deposits  of  the  “third  phase”  and  transgres-
sive Karpatian marine marls, indicate the possibility of an addi-
tional  marine  connection  between  the  Mediterranean  and/or
Indo-Pacific Ocean situated south-east of the  investigated area.

Acknowledgments:  The  authors  wish  to  thank  Professor
Ana Sokač for her comments and discussion about the ostra-
cod  fauna.  Professor  Vladimir  Bermanec  and  Hrvoje  Posi-
lović, B.Sc. from the Faculty of Science, Zagreb provided us
with  SEM  microphotographs.  The  authors  are  very  grateful
to Ivo Suša from the Croatian Geological Survey for techni-
cal support.

Appendix

New ostracod species

The following abbreviations are used in the description of

the new ostracod species: LV and RV – left and right
valves; H and L – height and length; n – number of mea-
sured specimens; and – female and male.

Class: Ostracoda Latreille, 1806

Order: Podocopida Sars, 1866

Suborder: Podocopina Sars, 1866

Superfamily: Cypridoidea Baird, 1845

Family: Candonidae Kaufmann, 1900

Subfamily: Candoninae Kaufmann, 1900

Genus: Fabaeformiscandona Krstić, 1972

Generic characters of this new species are according to

those  given  by  Van  Morkhoven  (1962).  Carapace  usually
elongated  in  lateral  view,  greatest  H  close  to  1/2  L.  Mostly
with significant sexual dimorphism. LV usually with poste-
ro-dorsal  lobe,  overlapping  the  RV  (diagnosis  after  Meisch
2000).

The genus Fabaeformiscandona comprises ca. 50 recent

species  (Meisch  2000).  The  oldest  fossil  representatives  of
this genus are known from the Early and Middle Miocene of
Europe  (Janz  1997;  Witt  1998,  2000;  Schäfer  2005).  Some
recent species of the genus Fabaeformiscandona  frequently
occur in Pliocene and Pleistocene (Meisch 2000).

Fabaeformiscandona slavonica nov. sp.

Fig. 3.1—7

M a t e r i a l : 46 valves.

♀

♂

258

HAJEK-TADESSE, BELAK, SREMAC, VRSALJKO and WACHA

E t y m o l o g y : Type locality is situated in Slavonia, re-

gion in eastern Croatia.

H o l o t y p e : Right valve, Fs-1, HGI-CGS Hr/Sa 2006/

HTV. Fig. 3.1.

T y p e - l e v e l : Lower Miocene/Upper Ottnangian—Lower

Karpatian, brackish deposits.

T y p e - l o c a l i t y : Sadovi, Mt Požeška gora, North Croat-

ian Basin.

P a r a t y p e s :

Left valve Fs-2, HGI-CGS Hr/Sa 2006/HTV. Fig. 3.2.
Right valve Fs-3, HGI-CGS Hr/Sa 2006/HTV. Fig. 3.5.
Left valve Fs-4, HGI-CGS Hr/Sa 2006/HTV. Fig. 3.4.

D i a g n o s i s : Fabaeformiscandona slavonica is best rec-

ognized in the external lateral view of right valve. The dorsal
margin of RV is specific with swale.

D e s c r i p t i o n : Valves smooth, strongly calcified, with

variable H/L ratio; greatest H situated distinctly behind mid—
length. Carapace laterally compressed in dorsal view. Dorsal
margin of the LV overlaps the RV. Sexual dimorphism present.

Male. Valves elongated less high than in the female. Dor-

sal margin slightly convex. Ventral margin slightly concave.
Anterior end rounded. Posterior end almost straight and ob-
liquely cut, forming a sharp angle with the postero-ventral
margin. LV (interior) slightly turned inwards in the upper
part of the posterior end. Males always present.

Female. RV with similar shape to male RV, slightly higher

than male. RV with swale on dorsal margin. LV with distinct
convex dorsal margin, gently and continuously sloping
down towards anterior and posterior ends (swale missing).
While anterior end is gently rounded, posterior end is almost
evenly cut off, forming a small point with the afferent ven-
tral margin. LV with maximum width in the central part.

M e a s u r e m e n t s (mm):

Holotype RV (Fs-1, HGI-CGS Hr/Sa 2006/HTV):

L = 1.04, H = 0.43.

Paratype LV (Fs-2, HGI-CGS Hr/Sa 2006/HTV):

L = 1.02, H = 0.38.

RV (Fs-3, HGI-CGS Hr/Sa 2006/HTV): L = 1.01,

H = 0.53.

LV (Fs-4, HGI-CGS Hr/Sa 2006/HTV): L = 1.02,

H = 0.54.

R a n g e s o f s i z e s o f s e l e c t e d s p e c i m e n s :

L (LV) = 1.03—0.99 mm (n=2); H (LV) = 0.39—

0.36 mm (n = 2).

L (LV) = 0.98 mm (n=1); H (LV) = 0.49 mm (n = 1).
L (RV) = 1.03—0.94 mm (n = 5); H (RV) = 0.41—

0.37 mm (n = 5).

L (RV) = 1.00—0.96 mm (n = 2); H (RV) = 0.51—

0.49 mm (n = 2).

R e m a r k s : The Fabaeformiscandona slavonica nov. sp.

has to be compared to the known described Miocene species:
Fabaeformiscandona fabaeformis (Fischer) and F. cf. bala-
tonica  (Daday)  (sensu  Janz  1997:  p. 3,  figs. 3—10;  p. 4,
figs. 1,  2);  F.  pokornyi  (Kheil)  (sensu  Witt  1998:  p. 1,
figs. 2—8;  2000:  p. 2,  fig. 3);  and  Fabaeformiscandona  sp.
(sensu Schäfer et al. 2005: p. 2, figs. 9—11), however, none is
close to the new species.

Other Fabaeformiscandona species have been investigated.

The new species differs from all of these. In general, Fabae-

formiscandona  slavonica  nov.  sp.  can  easily  be  distin-
guished  from  all  its  congeners  in  the  postero-dorsal  part  of
the male and female RV. Only the carapace of recent Fabae-
formiscandona  fabaeformis  (Fischer)  is  similar  in  dorsal
view.  Female  LVs  of  Fabaeformiscandona  slavonica    nov.
sp.,  resemble  those  of  Fabaeformiscandona  levanderi  (Hir-
schmann)  in  shape.  The  female  LVs  can  easily  be  distin-
guished from each other by the posterior and postero-ventral
part of the valve.

Family: Cyprididae Baird, 1845

Subfamily: Herpetocypridinae Kaufmann, 1900

Genus: Herpetocypris Brady & Norman, 1889

The new species belongs to the genus Herpetocypris. Car-

apace elongated in lateral view, about 1.5—2.6 mm long. The
height always less than half the length. Both ends are round-
ed. Dorsal margin weakly rounded to almost straight. Cara-
pace laterally moderately compressed. Fused valve zones
small, pore canals straight, marginal septae absent. LV over-
laps RV along the entire valve margin (diagnosis after Van
Morkhoven 1963; Meisch 2000).

Six certain and several dubious species have been record-

ed  from  Europe  (Meisch  2000).  Fossil  species  of  Herpeto-
cypris  are  known  from  the  Early  and  Middle  Miocene  of
Europe (Sokač 1979; Sokač & Krstić 1987; Jurišić-Polšak et
al.  1993;  Witt  1999,  2002).  The  fossil  records  for  some  re-
cent  species  of  genus  Herpetocypris  are  common  from  the
Pleistocene (Van Morkhoven 1963; Meisch 2000).

Herpetocypris sadovii nov. sp.

Fig. 4.1—3

M a t e r i a l : 15 valves.
E t y m o l o g y : After the type section in which it was first

found.

H o l o t y p e : Right valve, Hb-1, HGI-CGS Hr/Sa 2006/

HTV. Fig. 4.3.

T y p e - l e v e l : Lower Miocene/Upper Ottnangian—Lower

Karpatian, brackish deposits.

T y p e - l o c a l i t y : Sadovi, Mt Požeška gora, North Croat-

ian Basin.

P a r a t y p e : Left valve Hb-2, HGI-CGS Hr/Sa 2006/

HTV, sample Sa 10.

D i a g n o s i s : Distinctly elongated valve with rounded an-

terior and posterior end.

D e s c r i p t i o n : A large, elongate, well calcified valve.

Smooth surface. Dorsal margin almost straight in the middle
and sub parallel to the ventral line. Transitions from dorsal
margin to anterior and posterior ends are gradual. Ventral mar-
gin slightly concave in the middle. Anterior and posterior end
rounded, but anterior end broader than the posterior one.

Fused valve zones small. Marginal pore canals strait and

short. Muscle scars weakly evident. No distinct sexual di-
morphism could be detected in the examined material.

M e a s u r e m e n t s (mm):

Holotype RV Hb-1, HGI-CGS Hr/Sa 2006/HTV:

L = 1.34, H = 0.48.

♂

♀

♂

♀

261

EARLY MIOCENE OSTRACODS FROM THE SADOVI SECTION (CROATIA)

Paratype LV Hb-2, HGI-CGS Hr/Sa 2006/HTV:

L = 1.19, H = 0.46.

L (LV) = 1.21 mm (n = 1): H (LV) = 0.46 mm (n = 1).
L (RV) = 1.33—1.29 mm (n = 3): H (RV) = 0.47—0.42 mm

(n = 3).

R e m a r k s : Herpetocypris sadovii nov. sp. comes close

to Herpetocypris  chevreuxi (Sars). The two species differ in
the following characters: a) size –  H. chevreuxi are longer
than  H. sadovii; b) ventral margin – in   Herpetocypris  sa-
dovii  nov.  sp.  ventral  margin  is  more  concave;  c)  posterior
end  –  different  shape  in  postero-dorsal  part  of  valve.  The
new  species  Herpetocypris  sadovii  differs  from  the  Early
and  Middle  Miocene  Herpetocypris  species:  Herpetocypris
sp. A (sensu Witt 1999: p. 2, fig. 6—9; 2002: p. 2, fig. 5—8);
Herpetocypris  snegotini  (Krstić)  (sensu  Sokač  1979:  p. 2,
figs. 1,  3,  5);  Herpetocypris  sp.  (sensu  Jurišić-Polšak  et  al.
1993: p. 1, figs. 1, 6, 9, 12) in shape of valves, especially in
area of dorsal margin, and anterior/posterior ends.

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