GEOLOGICA CARPATHICA, 54, 1, BRATISLAVA, FEBRUARY 2003
9 — 20
THE OSTRACOD GENUS NIPPONOCYTHERE ISHIZAKI, 1971 FROM
THE MIDDLE MIOCENE OF THE FORE-CARPATHIAN DEPRESSION,
CENTRAL PARATETHYS; ITS ORIGIN AND PALEOENVIRONMENT
and GIUSEPPE AIELLO
Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warszawa, Poland; firstname.lastname@example.org
Dipartimento di Scienze della Terra, Università degli Studi di Napoli “Federico II” Largo S. Marcellino 10, Napoli, Italy
(Manuscript received April 4, 2002; accepted in revised form October 3, 2002)
Abstract: Two new ostracod species, belonging to the loxoconchid genus Nipponocythere Ishizaki, 1971, i.e. N. karsyensis
sp. nov. and N. silesiensis sp. nov., are described from the Middle Miocene (Badenian) of the Fore-Carpathian Depres-
sion, Central Paratethys. We consider them “exotic” forms, that is rare and previously unknown from the Badenian of the
Paratethys. Their occurrence in the Middle Miocene of the Polish part of the Central Paratethys is thought to prove
oceanic water influence into the northernmost parts of the Fore-Carpathian Depression (basins). The unusual morpho-
metric features of these Nipponocythere species and of their associated “exotic” ostracods, in the Upper Badenian of the
Upper Silesia, is presumed to be due to a paleoenvironment rich in food and undersaturated in CaCO
Key words: Middle Miocene, Fore-Carpathian Depression, paleoecology, paleogeography, Ostracoda.
Among the Middle Miocene (Badenian) ostracods of the Pol-
ish part of the Fore-Carpathian Depression (Central Parat-
ethys), are two species of Nipponocythere Ishizaki, 1971,
which was unknown previously from Europe (except for the
Basque Basin, i.e. its Atlantic border), including the whole
Mediterranean, prior to the Upper Miocene. These species are
herein described as Nipponocythere karsyensis sp. nov., from
the Lower Badenian, and Nipponocythere silesiensis sp. nov.
from the Upper Badenian of the studied area.
Among the Middle Miocene ostracods of the Central Para-
tethys, Nipponocythere are “exotic” forms, that is rare and so
far unknown from the Badenian of Paratethys, although occa-
sionally encountered in past and Recent oceans. Other exot-
ics, such as Cluthia, Xylocythere, Nunana and Elofsonia were
described earlier (Szczechura 1986, 1995; Aiello & Szczechu-
ra 2001a, 2002), while the discovery of others will be pub-
lished soon. The presence of Nipponocythere in the Middle
Miocene (Lower and Upper Badenian) of the northern part of
the Central Paratethys provides additional data for the recon-
struction of the Neogene (Middle Miocene) paleogeography
and paleoecology of this area.
The material described here is housed at the Institute of Pa-
leobiology of the Polish Academy of Sciences in Warszawa
Material (General setting)
In the Fore-Carpathian Depression of southern Poland,
which represents the northern margin of the Central Para-
tethys, ostracods belonging to Nipponocythere Ishizaki, 1971,
occur throughout the entire Middle Miocene (Badenian). The
representatives of this genus were found in the Badenian sedi-
ments of the Działoszyce Trough (Posądza 10-S borehole) as
well as in the Korytnica Basin (Karsy outcrop), on the south-
ern slopes of the Holy Cross Mts, and in the Upper Silesia
(Gliwice G-19 and Gliwice G-21 boreholes) (Fig. 1).
Posądza 10-S borehole (Działoszyce Trough)
A species of Nipponocythere was found in the Lower Bad-
enian (corresponding to the Langhian stage) marly sedi-
ments in the borehole Posądza 10-S (depth 101.8—100.2 m),
in the Działoszyce Trough (Osmólski 1972; Szczechura
2000). The entire section, representing Badenian and Sarma-
tian evaporite-bearing deposits, is about 90 m thick. It over-
lies Upper Cretaceous carbonates and consists of marls and
marly clays passing upwards into clays. Ostracods are asso-
ciated with rich foraminiferal assemblages (both planktonic
and benthic) indicating a bathyal and rather oligotrophic en-
vironment with warm surface waters (Szczechura 2000). The
chronology was based on planktonic foraminifers (mostly
Praeorbulina and Orbulina). Bolboforms (calcareous phy-
toplankton) and radiolarians are very rare. Ostracods (men-
tioned in Szczechura 2000) are also scarce and mostly poor-
ly preserved; they include species of Argilloecia, Buntonia,
Bythocypris, Cnestocythere, Costa, Cytherella, Cytheropter-
on, Eucythere, ?Eucytherura, Henryhowella, Krithe, Paijen-
borchella, Parakrithe, Pterygocythereis, Saida and Xestole-
beris. These genera are mostly typical of the deeper waters
and rather ubiquitous in their geographical (and paleogeo-
graphical) distribution. Both the taxonomic composition of
the ostracod assemblage and their rarity confirm the bathy-
metrical (and perhaps trophic) conditions suggested by the
10 SZCZECHURA and AIELLO
Karsy outcrop (Korytnica Basin)
Nipponocythere was also found in the Lower Badenian silty
sediments, in the so called Korytnica Clays, cropping out in
Karsy near Korytnica (Bałuk & Radwański 1977). The Koryt-
nica Clays are represented by 40—60 m of undisturbed and
highly fossiliferous marly silts and clays, overlain by marly
sands and red-algal (lithothamnian) limestones. Nippono-
cythere is associated with an unusually abundant, diverse and
very well preserved microfauna consisting mostly of benthic
and planktonic foraminifers and ostracods, although, rare ra-
diolarians and bolboforms are also present. Calcareous benth-
ic foraminifers are dominated by forms typical of shallow
(shelf) waters. The presence of large foraminifers (Borelis,
Amphistegina) and some characteristic planktonic foramini-
fers (e.g. Globigerinoides, Globoquadrina, Orbulina) sug-
gests a warm-water environment (cf. Rögl & Brandstötter
1993; the present authors’ observations). The planktonic as
well as larger foraminifers were used as biostratigraphical
The ostracod assemblage consists of abundant, well pre-
served, adult and juveniles; mostly the ubiquitous taxa Acan-
thocythereis, Aurila, Bosquetina, Buntonia (B. subulata subu-
lata), Callistocythere, Costa, Cytherella, Cytheridea, Cyther-
opteron, Hemicytherura, Incongruellina, Loxoconcha, Ptery-
gocythereis, Semicytherura and Xestoleberis. These taxa indi-
cate a low-energy near-shore (?infralittoral) environment,
most probably with plants. Rare specimens belonging to the
genera Henryhowella, Krithe, Paijenborchella and Parakrithe
and species known to prefer deeper-water conditions such as
Buntonia dertonensis, may suggest influence of the deeper
waters, possibly by means of bottom currents.
It should be noted, however, that the Korytnica Clays fau-
nas are strongly variable, both laterally and vertically, so the
studied samples may actually represent mixtures of ostracods
of somewhat different origin. According to Radwańska
(1992), working on the fish otoliths from the Korytnica Clays,
there are mixed assemblages, containing both shallow and
deeper-waters taxa, which could be the result of storm activi-
ty. Radwańska (op. cit.) proposed that the depth of deposition
of the Korytnica Clays corresponded to the littoral zone.
Kowalewski & Miśniakiewicz (1993), working on the shell
material from the Korytnica Clays, considered it a storm de-
Gliwice G-19 and G-21 boreholes (Działoszyce
Nipponocythere was also recorded from Upper Badenian
(corresponding to the Serravallian stage of the Mediterranean
region) argillaceous sediments, with limestones and tuff inter-
calations, about 70 m thick, overlying evaporites, in the Gli-
wice G-19 (depth 51.0—90.0 m) and G-21 (depth 19.2—
51.0 m) boreholes (Gonera 1997; Alexandrowicz 1997;
Szczechura 1997). The entire section consists of nearly 290 m
thick Badenian sediments, overlying Carboniferous strata and
covered by Quaternary deposits. The associated microfossils
include very diverse, abundant, and well preserved fora-
minifers (Alexandrowicz 1963, 1997; Gonera 1997), radio-
larians (Barwicz-Piskorz 1997), bolboforms (Szczechura
1997), dinoflagellates (Gedl 1997), holothurian sclerites (Gór-
ka 1997), pteropods (Janssen & Zorn 1993) and diatoms (Wit-
kowski & Gonera 1997). Interestingly, some microfossils are
dark and/or infilled with pyrite. Mainly microfossils were
used for the biostratigraphical subdivision of the studied sec-
tion, which has been attributed by Peryt (1997) to the calcare-
ous nannoplankton zone NN6. The foraminifers belong to a
so-called “IIIB assemblage” (Gonera 1997), distinguished by
Alexandrowicz (1963) and recognized by him from numerous
Upper Badenian sections of the Fore-Carpathian Depression,
although exhibiting much variation both geographically and
stratigraphically. The abundant benthic (mostly calcareous)
foraminifers include epifaunal (including epiphytics) as well
as infaunal elements. Planktonic forms are less diversified and
Witkowski & Gonera (1997) state that the diatoms, consist-
ing mostly of benthic (epiphytic and sediment dwelling)
forms, indicate deposition within the photic zone, in a rela-
tively shallow, quiet-water environment. It seems important,
that Thalassionema nitzschioides (although rare) occurs
among the diatom flora recognized by Witkowski & Gonera
(1997). This species was regarded as an upwelling indicator in
the Miocene of the eastern equatorial Pacific (Funnell et al.
1996). On the other hand, Barwicz-Piskorz (1997) considered
the radiolarian bloom in the studied part of the section from
Gliwice as an effect of volcanic activity, which enriched sea-
water in silica. At the same time Barwicz-Piskorz (1997) also
pointed out that the occurrence of Nasselaria is characteristic
of upwelling areas. Moreover, dinoflagellates (as other phy-
toplankton) also seem to prefer zones of upwelling (Brasier
1995; Hutchings et al. 1995; Summerhayes et al. 1995). The
microfossils therefore suggest an especially attractive, mostly
eutrophic, open marine (oceanic origin), but not very deep
(?circalitoral) environment, most probably influenced by up-
It should be added that some authors (e.g. Łuczkowska
1974; Szczechura 1982, 2000; Durakiewicz et al. 1997; Gon-
era 1997) suggested that during the late Badenian the climate
in the Fore-Carpathian Depression area was cooler than that
during the early Badenian, that is there was drop in tempera-
ture before the deposition of evaporites.
The ostracods are represented by numerous adults and juve-
niles (mostly disarticulated valves), and despite being small-
sized and thin-shelled they are, in general, well preserved sug-
gesting quiet-water sedimentation. The species richness (more
than one hundred species) as well as the abundance of ostra-
cods seems to suggest particularly favourable environmental
conditions. The occurrence of phytal, shallow-water forms ap-
pears to demonstrate shallow, photic zone depths. The ostra-
cod assemblage contains quite common representatives of
Argilloecia, Callistocythere, Cytherois, Cytheropteron, Hemi-
cytherura, Loxoconcha, Microcytherura, Paracytherois,
Pseudocytherura, Sagmatocythere, Semicytherura, Xestoleb-
eris, while species belonging to the genera Henryhowella,
Krithe, Parakrithe and Pseudocythere are less common. Par-
ticularly significant, however, is the cooccurrence of ostra-
cods (besides Nipponocythere and those earlier mentioned
Cluthia, Xylocythere, Elofsonia and Nunana) previously un-
known in the Middle Miocene of the Central Paratethys. They
THE OSTRACOD GENUS NIPPONOCYTHERE ISHIZAKI, 1971 11
include species of Rimacytheropteron, Swainocythere, Kuni-
hirella, Loxoconchidea, Polycope, Saida, Hemipara-
cytheridea and Ruggieriella. All these forms are mentioned,
among ostracods of different age (mostly Cainozoic) and dif-
ferent areas (including Recent oceans), as typical of deeper
waters (Whatley & Ayress 1988; Coles et al. 1990; Van
Harten 1990; Rodriguez-Lazaro & Garcia-Zarraga 1996). In
the Mediterranean basins, the above mentioned genera first
appeared (as is so far known) in the Upper Miocene. From our
prior knowledge of the ostracod assemblages of the Middle
Miocene of the Fore-Carpathian Depression, the present oc-
currence of these “exotic” taxa is unique.
It is worth mentioning, that in the supra-evaporitic sedi-
ments in the Gliwice sections, the diversity of ostracods grad-
ually increases (suggesting gradual appearance of new species
by migration) and reaches a maximum at the height of the late
Badenian transgression in the studied area.
Discussion and conclusions
According to a review by Barra (1995) the genus Nippono-
cythere is represented by more than 20 species and first ap-
peared in Costa Rica, where it spans the Lower and basal
Middle Miocene. During the Middle Miocene it entered the
Western Atlantic including central Haiti, Trinidad and Vene-
zuela. Nipponocythere persisted in the Caribbean area until
the Recent, but during the Pliocene it invaded the southwest-
ern Pacific and South China Sea, while during the Pleistocene
it appeared in southwestern Taiwan and central Japan. In the
Recent it also colonized the northwestern Pacific, including
Japan and the East China Sea, and the Gulf of Panama.
If (as is suggested here) Nipponocythere is congeneric with
Bidgeecythere McKenzie, Reyment et Reyment, 1993, its first
appearance could in fact have been in southwestern Australia
(Victoria), where Bidgeecythere ranges from Eocene to Mid-
dle Miocene (McKenzie 1974; McKenzie et al. 1993). It sub-
sequently spread to the Caribbean region; however, the repre-
Nipponocythere) are also recorded from the Lower Eocene of
the Atlantic border of the Basque Basin (Biscay Synclinori-
um) (Rodriguez-Lazaro & Garcia Zarraga 1996). It is worth
mentioning that other ostracods associated with Nippono-
cythere in the Middle Miocene of the Central Paratethys, for
example Cluthia (Szczechura 1986; Ayress & Drapala 1996)
and Nunana (Aiello & Szczechura 2001a) appeared earlier (in
the Eocene) in the Southern Hemisphere than in the Northern
The existence of Nipponocythere in both hemispheres dur-
ing the Eocene provides new evidence of the possibilities of
exchange of this microfaunal group (ostracods) between these
areas, perhaps along the already existing latitudinal routes of
migration (Benson 1990; Coles et al. 1990).
The presence of Nipponocythere in the Atlantic (and the
Central Paratethys) during the Middle Miocene suggests that
it arrived in the Fore-Carpathian Depression at the same time.
Fig. 1. Extent of the Middle Miocene sea in the Fore-Carpathian Depression and the location of sampled sections. 1 – edge line of the Car-
pathian nappes. 2 – northern edge of the Miocene Basin. 3 – extent of the continuous cover of the terrestrial “Poznań series”. 4 – Mi-
ocene deposits from the Fore-Carpathian Depression in Poland. 5 – locality (1 – Posądza, 2 – Karsy, 3 – Gliwice, 4 – Brno). 6 – cit-
ies. (Modified after Łuczkowska 1998.)
12 SZCZECHURA and AIELLO
It is not impossible, however, that Nipponocythere (like other
ostracods known from the Middle Miocene of the northern
part of the Central Paratethys) could already have invaded the
more southern part of the Central Paratethys, at least during
the Early Miocene. The presence of Nipponocythere in the
Lower Badenian of Moravia (Fig. 1) suggests its northward
migration from southern to central Europe, possibly through
the Moravian Gate. The existence of an early Middle Miocene
connection between the Atlantic Ocean and the Mediterranean
basins, via the Iberian Portal, was suggested (using ostracod
distributions) by Benson (1976) as well as by Nascimento &
Říha (1996). These latter authors found more than thirty ostra-
cod species common to the Middle Miocene of Portugal and
the Central Paratethys as well as the Mediterranean Basin.
Szczechura (1994, 1997, 2000) documented the existence of
the early Middle Miocene extension of the Atlantic influence
into the Central Paratethys, using ostracods as well as bolbo-
forms and foraminifers. This early Middle Miocene connec-
tion between Atlantic and the Central Paratethys basins, via
the Mediterranean, has also been confirmed and/or suggested
by other fossil groups, such as the pteropods (Janssen & Zorn
1993), bivalves (Studencka et al. 1998) and fish otoliths (Rad-
wańska 1992). However, according to Radwańska (1992),
Rögl (1998) and others there was an Indo-Pacific influence
during the Badenian (mostly the Early Badenian), in the Cen-
The occurrence of Nipponocythere (although belonging to
different species) in the Lower as well as in the Upper Bade-
nian of the Fore-Carpathian Depression indicates that this ge-
nus could have persisted there through the Middle Badenian
salinity crisis, like numerous other associated ostracod genera
and species. Refugia must have existed somewhere, or Nip-
ponocythere silesiensis reappeared in the Upper Badenian of
the Upper Silesia as a result of a new (or only enlarged) Atlan-
tic oceanic influence in the Central Paratethys. This latter sup-
position may be confirmed by the invasion, during the Late
Badenian in the Polish part of the Fore-Carpathian Depres-
sion, of a new ostracod fauna, which has not hitherto been re-
corded in the sub-evaporitic sediments (Szczechura 1995,
1996, 1998; Aiello & Szczechura 2001a, 2002). The late Mid-
dle Miocene distribution of Carinocythereis in Europe (Szcze-
chura 1998; Aiello & Szczechura 2001b) indicates beyond
doubt that, at that time, a connection existed between the
northern part of the Central Paratethys and the Mediterranean
basins, possibly via the southeastern areas of Europe. At the
same time, other common groups of microfossils appeared in-
cluding bolboforms (Szczechura 1997), radiolarians (Bar-
wicz-Piskorz 1997) and pteropods (Janssen & Zorn 1993). In-
terestingly, some of them are also known from the northern
The literature (e.g. Ishizaki 1981; van den Bold 1985;
Whatley & Ayress 1988; Coles et al. 1990; Van Harten 1990;
McKenzie et al. 1993; Bonaduce et al. 1994; Barra 1995) in-
dicates, that Nipponocythere is a marine genus with no partic-
ular environmental preferences in respect of temperature and
depth. Its Recent and fossil representatives are recorded from
shallow (though generally not very shallow) as well as deep
waters in various areas of both hemispheres, independently of
The distribution of Nipponocythere in the Middle Miocene
of the Fore-Carpathian Depression seems to provide, howev-
er, some new indications on its environmental preferences.
In the Middle Miocene of the Fore-Carpathian Depression
Nipponocythere occurs, most probably in situ, in the lower
Middle Miocene sediments from the Działoszyce Trough in
Posądza (Szczechura 2000), where it is very rare and is asso-
ciated with taxa typical of the normal marine, bathyal environ-
ments, characterized by cold and rather oligothrophic bottom
waters, as in the Lower Badenian of the Czech Republic (Říha
1989). In the approximately coeval but shallow-water sedi-
ments of the Korytnica Basin (Karsy), Nipponocythere could
be derived (by storm activity), as are some other coexisting
ostracod species and this, therefore, may be not have been its
In the Upper Badenian of Upper Silesia (Gliwice area) Nip-
ponocythere is more common than in the Korytnica Basin and
it certainly proliferated there. However, the environment
seems to have been shallow marine, probably nearshore and,
as is suggested by the associated microfauna, rich in food sup-
ply. Even more surprising is that Nipponocythere is accompa-
nied by deeper-water ostracods and the newly described, “ex-
otic” species. It also seems significant that all of them are
generally small and thin-shelled, and sometimes dark in co-
lour. Large and thick-shelled forms are lacking. Moreover,
this ostracod assemblage seems to be unique when compared
to others known from the Middle Miocene deposits of the
According to Szczechura (1986), the first appearance in the
Upper Badenian of the Central Paratethys of Cluthia mioceni-
ca (one of the “exotic” forms from Gliwice), was the result of
the cooling; Cluthia is known as a northern “guest”, that is a
cold-water ostracod form (cf. e.g. Carbonnel & Ballesio
1982). Also Brouwers et al. (2000), reviewing the spatiotem-
poral distribution of Cluthia, have concluded that this genus
followed its preferred lower temperature range, invading shal-
low but cold waters. A similar relationship, that is the migra-
tion of Recent deep-water ostracods onto the shelf (at shallow
depths), in the southeastern Pacific, as the result of cold tem-
perature and low salinity, was suggested by Whatley et al.
(1997); these authors, however, highlighted the importance of
upwelling for ostracod distribution in such circumstances as
the southern end of the Magellan Strait.
These conclusions seem to be supported by the case from
Gliwice, where ostracods thrived in rather shallow (relatively
shallow) but cool waters influenced by upwelling. Such up-
welling may explain the high diversity and abundance of os-
tracods; but it does not explain the distinctive taxonomic
structure or the peculiar morphometry of the “exotic” as well
as the associated ostracods. It is broadly known (e.g. Murray
1991, 1995; Brasier 1995; Dingle 1995), that the high produc-
tivity of the surface waters may have an impact on bottom-
water chemistry and, in consequence, the distribution as well
as the character of the fauna (including ostracods).
In this respect it seems significant that the ostracod assem-
blage that is most taxonomically similar to that from Gliwice
THE OSTRACOD GENUS NIPPONOCYTHERE ISHIZAKI, 1971 13
(containing numerous common genera and some conspecific
and/or very close species), is described from the Upper Mi-
ocene of Italy (Piedmont, Sicily, Calabria) (Ciampo 1986) and
from the Pleistocene of Calabria (Colalongo & Pasini 1980).
In the Pleistocene of Calabria (op. cit.) ostracods occur in sed-
iments, which according to the cited authors were deposited
under poorly aerated, bathyal conditions. Elements of the Gli-
wice fauna discussed here also occur in the Azores at the Mid-
Atlantic Ridge (Van Harten 1990), as well as in the Scoresby
Sund Fiord of East Greenland (Whatley et al. 1996). This lat-
ter site represents a fairly deep-water basin, silled in its front
and interpreted as a stagnating basin rich in organic carbon
and at least seasonally impoverished in oxygen in its bottom
waters. The ostracods (somewhat) similar to those from Gli-
wice are also described by Coles et al. (1996) from the Qua-
ternary carbonate mounds, associated with gas seepages, from
the Porcupine Basin (offshore western Ireland). This rather
deep-water ostracod association (mostly paradoxostomatids)
contains xylophile (although without Xylocythere) forms and
is thought to be an indirect result of nutrient increase. The
large diversity and abundance of ostracods is correlated here
with wide niche variety. Interestingly, one of the “exotic”
forms from Gliwice, the genus Xylocythere, is known as the
typical xylophile (wood-loving) and at the same time deep-
water form (Maddocks & Steineck 1987; Steineck et al.
1990). Van Harten (1992) found it in deep-water sediments, in
the Pacific, coexisting with ostracods characteristic for shal-
low waters, associated with hydrothermal vents. He consid-
ered that it proliferated in the highly eutrophic (and specifical-
ly, rich in chemical energy) environment, impoverished in
oxygen (Van Harten 1992, 1993). According to Van Harten
(op. cit.), this latter factor may have caused the poor biocalci-
fication of the ostracod carapaces, expressed in their thin
In almost all the cases reviewed above, the ostracods dis-
cussed are correlated with an environment rich in food and
poorly oxygenated. Therefore, both these environmental fac-
tors could apply to the Upper Badenian bottom waters in Up-
per Silesia. This suggestion may be additionally supported by
the occurrence of pyrite as well as the dark colour of the mi-
crofossils (including ostracods) (cf. Neale 1971; Oertli 1971).
According to Oertli (1971), the small size and pyritization of
ostracod shells, is a consequence of reducing environments
rich in organic carbon. Neale (1971), working on the Lower
Cretaceous microfauna (foraminifers and ostracods) of the
Speeton Clay, England, suggested that pH, food supply and
rate of sedimentation are the most important factors influenc-
ing the abundance and composition of faunas. Moreover,
Neale (op. cit.) stated that at least partial de-oxygenation of
the bottom waters was tolerated only by the specialized fauna,
represented by small forms, as opposed to the large and varied
forms occurring in horizons abundant in food and CaCO
A close relationship between the availability of food and
the chemistry (mostly pH and CO
content) of the bottom wa-
ters, as influencing biocalcification in ostracods, have also
been considered by Peypouquet (1977), Carbonel & Hoibian
(1988), Peypouquet et al. (1988), Babinot et al. (1991) and
Braccini & Peypouquet (1996). Moreover, as is shown e.g. by
Curry (1999), Mezquita et al. (1999) and Cronin et al. (1999),
various (different) ostracods species and/or groups have their
particular ecological requiements, including physical and
It is worth mentioning, that a much more severe (mostly in
terms of the lowered oxygen content) environment was recog-
nized in the more central part of the Fore-Carpathian Depres-
sion, in the Jamnica borehole, near Tarnobrzeg (Szczechura
1995). In this section a few specimens of Xylocythere (X. car-
pathica) were found in association with a low diversity micro-
fauna (foraminifers and ostracods) and they were thought to
occur there in a stressed (low-oxygen) but at the same time
rather shallow-water environment. More recently, Xylocythere
carpathica was also recorded from the approximately coeval
sediments of the Działoszyce Trough (the Posądza 10-S bore-
hole) (Szczechura 2000). In the Posądza section, just above
the evaporites, benthic microfauna is absent, although radi-
olarians are abundant and seem to mark a distinct biosiliceous
event. Somewhat above, in the overlying sediments rather rare
and weakly diversified benthic and planktonic foraminifers
appear, indicating a stressed bottom water environment. They
are associated with a bispecific ostracod assemblage, consist-
ing of numerous (considering the small size of sample), un-
usually delicate and small-sized, adult and juvenile representa-
tives of Xylocythere (X. carpathica) and ?Microxestoleberis sp.
These findings (from Gliwice, Jamnica and Posądza) prove
that during the Later Badenian of the Fore-Carpathian Depres-
sion, the environment was more or less depleted in oxygen
and highly euthrophic, which favoured the development of
distinctive ostracod populations. In all these cases, abundance
of nutrients, dysaerobic bottom microhabitats (i.e. lowered pH
of the bottom waters), resulting in their undersaturation in
and causing poor biocalcification, allowed only small,
thin-shelled ostracods to proliferate.
Interestingly, small and thin-walled tests of calcareous
benthic foraminifers (mostly particular morphogroups) are
known from low-oxygen bottom water environments (Sen
Gupta & Machain-Castillo 1993; Gooday 1994; Kaiho 1994,
1999) resulting from high primary productivity of the surface
waters, and the consequent flux of organic matter to the sedi-
ment. These authors argued that biomineralization was more
difficult in the suboxic environment.
It must be added that the Upper Badenian marine environ-
ment of the Fore-Carpathian Depression was particularly sus-
ceptible to eutrophication. It could have been caused, except
for the (local) pericoastal upwelling, by contemporary volca-
nic activity enhancing primary productivity. In Upper Silesia
(Gliwice area), eutrophication could have been additionally
strengthened by its paleogeographical situation, that is its
marginal location within the Fore-Carpathian Depression en-
hancing support of the continentally derived organic matter,
sea level rise, as well as the existence of the so called Cracow
Ridge (Ney et al. 1974), a submarine barrier preventing ex-
change of waters between the Upper Silesia area and the rest
of the Fore-Carpathian Depression (an “inflow” circulation
model); the presence of some common species, in the western
as well as in the eastern part of the Polish part of the Fore-Car-
pathian Depression, indicates that there was water mass ex-
change (at least to some extent) between these areas. Another
factor favouring euthrophic conditions, resulting in the dys-
14 SZCZECHURA and AIELLO
THE OSTRACOD GENUS NIPPONOCYTHERE ISHIZAKI, 1971 15
aerobic bottom environment, was probably slow sedimenta-
tion (quiet environment, weak circulation), suggested by the
thin-shelled but well preserved ostracod remnants, only as
valves, and the vegetation (sea grasses) covering the seafloor.
At the same time the activity of vents and/or seeps in the Fore-
Carpathian Depression is not excluded.
According to Garlicki & Szybist (2001), working on the or-
ganic matter content in the Middle Miocene sediments from
the southern areas of the Polish part of the Fore-Carpathian
Depression south-west of Tarnów (Fig. 1), organic matter oc-
curs throughout the entire evaporites-bearing sections. In the
sediments overlying the evaporites it attains up to 9.3 % of
weight. Garlicki & Szybist (2001) considered these results
mostly the effect of high phytoplankton productivity. They
also found a close relationship between the supply of the or-
ganic matter and the bottom water conditions.
The following abbreviations are used: a – adult, C –
carapace, RV – right valve, LV – left valve.
Suborder Podocopina Sars, 1866
Superfamily Cytheracea Baird, 1850
Family Loxoconchidae Sars, 1925
The family assignment of the examined genus is far from
equivocal. Heinia was referred by its author (van den Bold
1985) to Cytheridae Baird, 1850; Bidgeecythere was consid-
ered by McKenzie et al. (1993) as belonging to Pecto-
cytheridae Hanai, 1957, while Nipponocythere was included
by Ishizaki (1971) as well as by Malz (1981) and Barra (1995)
in the Loxoconchidae Sars, 1925.
Genus Nipponocythere Ishizaki, 1971
We agree with Drapala & Ayress (1993), Bonaduce et al.
(1994) and Barra (1995) that Nipponocythere Ishizaki, 1971 is
congeneric with Heinia van den Bold, 1985. The genus Bid-
geecythere, described by McKenzie, Reyment et Reyment
(1993) from the Eocene of Australia, is considered here as a
junior synonym of Nipponocythere because it possesses all
Fig. 2. 1 – Nipponocythere karsyensis sp. nov., aLV, holotype;
a – lateral view, b – oblique ventral view, c – dorsal view;
ZPAL O.48/319. 2 – Nipponocythere karsyensis sp. nov., aRV;
a – lateral view, b – ventral view; ZPAL O.48/304. 3 – Nip-
ponocythere karsyensis sp. nov., aLV, paratype, ZPAL O.48/318.
4 – Nipponocythere karsyensis sp. nov., aRV, paratype; a – lat-
eral view, b – ventral view; ZPAL O.48/305. 5 – Nippono-
cythere karsyensis sp. nov., aC, right side, ZPAL V.27/731. 6 –
Nipponocythere silesiensis sp. nov., aRV; a – general internal
view, b – enlarged proximal part of the hinge margin, c – en-
larged distal part of the hinge margin; ZPAL O.48/20.
1—4 – Lower Badenian, Karsy outcrop; 5 – Lower Badenian,
Posądza 10-S borehole, depth 100.2 m; 6 – Upper Badenian, Gli-
wice G-19 borehole, depth 51.0 m. Scale bars = 1 mm.
the external and internal diagnostic features typical of the
above mentioned genus.
Nipponocythere silesiensis sp. nov.
(Fig. 2.6; Fig. 3.1—8)
H o l o t y p e : RV ZPAL O.48/60 illustrated in Fig. 3.4.
P a r a t y p e s : The specimens ZPAL O.48/62,61, illustrated
in Fig. 3.3,7.
T y p e h o r i z o n : Upper Badenian (upper part of the Mid-
T y p e l o c a l i t y : Gliwice (Upper Silesia).
D e r i v a t i o n o f t h e n a m e : silesiensis, occurring in
D i a g n o s i s : A Nipponocythere species rather weakly and
mostly centrally punctate, indistinctly (finely) ribbed an-
teroventrally, weakly reticulate postero-laterally, bearing a
short ear-shaped rib along the posterior part of the dorsal
M a t e r i a l : 15 adult valves and one adult carapace, well
D e s c r i p t i o n : Carapace small, subovate to subtriangular
in lateral view, weakly and almost evenly inflated laterally,
elongated and pointed posteriorly especially in the poster-
oventral part of RV. It is somewhat truncated in LV, being
moderately rounded anteriorly, straight dorsally, weakly in-
cised ventrally. The valve surface rather densely punctate,
mostly centrally; if well preserved provided with the indis-
tinct net work-like reticulation at the posterior end. A few
weak, subparallel ribs extend over the ventro-lateral surface.
The anterior margin is bordered by a more or less distinct but
rarely preserved, delicate frill-like flange. A short ear-
shaped rib occurs along and near the posterior part of the
Inner lamella moderately wide anteriorly and posteriorly. It
is subparallel to the valve margins. Vestibulum small, below
mid-height, in the anterior part, less distinct posteriorly. Mus-
cle scars (Fig. 3.8b) consist of four oval scars of the main
group and a thickly arcuate concave forward antennal scar.
Hinge (Fig. 2.6b—c; Fig. 3.8a) modified gongylodont. In the
RV the anterior terminal element is a locule which surrounds
the antero-median element; the median element is a long,
smooth groove with an antero-median boss-like tooth. The
postero-terminal element is a boss-like tooth. Complementary
elements occur in the LV hinge. Marginal pore canals simple,
rather rare; 7—9 in frontal part, while 6—7 in distal part.
V a r i a b i l i t y : It is expressed in size, and degree and ex-
tent of ornamentation.
D i m e n s i o n s o f a d u l t s ( i n m m ) :
RV LV LV
R e m a r k s : This species is similar to that described by
Drapala & Ayress (1993), from the Late Quaternary of Aus-
16 SZCZECHURA and AIELLO
THE OSTRACOD GENUS NIPPONOCYTHERE ISHIZAKI, 1971 17
tralia, as Nipponocythere colalongoae (Ciampo, 1985), pro-
posed by Bonaduce et al. (1994) as Nipponocythere
drapalaensis sp. nov. Nipponocythere silesiensis is more
elongate, more pointed posteriorly, more densely foveolate,
and bears a shorter, less pronounced ear-shaped rib along the
posterior part of the dorsal margin. In comparison with Nip-
ponocythere karsyensis sp. nov. from the Lower Badenian of
the Carpathian Foredeep, this species is more posteriorly elon-
gated, more laterally inflated, less densely and mostly central-
ly punctate, and only weakly reticulated at the posterior mar-
gin. Its posterodorsal rib is short, ear-shaped, and does not
D i s t r i b u t i o n : Upper Badenian of Upper Silesia, Gli-
wice (Central Paratethys).
Nipponocythere karsyensis sp. nov.
1983 Bosquetina carinella (Reuss, 1850); Říha, fig. 5, pl. 1, fig. 5.
H o l o t y p e : RV ZPAL O.48/319, illustrated in Fig. 2.1.
P a r a t y p e s : The specimens ZPAL O.48/318, 305, illus-
trated in Fig. 2.3,4, respectively.
T y p e h o r i z o n : Lower Badenian (lower part of the Mid-
T y p e l o c a l i t y : Karsy (Korytnica Bay).
D e r i v a t i o n o f t h e n a m e : karsyensis, occurring in
D i a g n o s i s : A Nipponocythere species being densely and
rather evenly punctate, mostly centrally (except the anterior
and posterior ends), and somewhat reticulate posteriorly.
Along the posterior part of the dorsal margin there is a short,
anteriorly branching rib.
M a t e r i a l : Seven adult valves and two carapaces, well
D e s c r i p t i o n : Small, subcuneate in lateral outline, weak-
ly and almost evenly inflated laterally, broadly rounded ante-
riorly, distinctly elongated and weakly truncated posteriorly,
nearly straight dorsally, indistincly incised ventrally. The
valve surface somewhat rough, densely and rather regularly
punctate, mostly centrally, with a weak reticulation in the pos-
terior part. Indistinct subparallel ribs cover ventro-lateral sur-
face, while a short but pronounced rib, branching frontally,
occurs along the posterior part of the dorsal margin. Anterior
margin bordered by a delicate frill-like flange.
Inner lamella moderately wide, with small anterior vestibu-
lum. Muscle scars consist of four semicircular adductors ar-
ranged in a vertical row, and one, somewhat heart-shaped
frontal scar. Hinge gongylodont, in the right valve consisting
of the anterior tooth, bounded by a narrow furrow, which
passes into the median groove, and posteriorly into a steep
dentate bar passing into terminal knob-like tooth. Marginal
pore canals not visible.
V a r i a b i l i t y : This involves shape, especially lateral out-
line, and posterodorsal inflation, probably resulting from sex-
ual dimorphism. Supposed males seem to be more triangular
in lateral outline, higher anteriorly and less posterodorsally in-
flated than supposed females. However the sexual dimor-
phism found in Nipponocythere drapalaensis pithekoussaii
(Barra, 1995), from the Late Quaternary of the Mediterranean
Basin (Isle of Ischia) is restricted mostly to a different lenght
to height ratio of males and females. The type of punctation of
specimens referred to N. silesiensis is somewhat variable and
may be a consequence of different states of preservation.
D i m e n s i o n s o f a d u l t s ( i n m m ) :
LV LV RV
Fig. 3. 1 – Nipponocythere silesiensis sp. nov., aLV, lateral view,
ZPAL O.48/555. 2 – Nipponocythere silesiensis sp. nov., aLV, lat-
eral view, ZPAL O.48/556. 3 – Nipponocythere silesiensis sp.
nov., aLV, paratype, lateral view, ZPAL O.48/62. 4 – Nippono-
cythere silesiensis sp. nov., aRV, holotype, lateral view, ZPAL
O.48/60. 5 – Nipponocythere silesiensis sp. nov., aLV; a – lateral
view, b – dorsal view; ZPAL O.48/294. 6 – Nipponocythere
silesiensis sp. nov., aRV; a – dorsal view, b – ventral view, c –
lateral view, d – enlarged details of the anterior margin; ZPAL
O.48/23. 7 – Nipponocythere silesiensis sp. nov., aLV, paratype,
internal view, ZPAL O.48/61. 8 – Nipponocythere silesiensis sp.
nov., aRV; a – general internal view, b – enlarged muscle scars
field; ZPAL O.48/554. 1, 3—5, 7, 8 – Upper Badenian, Gliwice
G-21 borehole, depth 40.10 m; 2 – Upper Badenian, Gliwice G-21
borehole, depth 51.0 m; 6 – Upper Badenian, Gliwice G-19 bore-
hole, depth 51.0 m. Scale bars = 0.1 mm.
R e m a r k s : In comparison with Nipponocythere parva
(Colalongo et Pasini, 1980) from the Plio-Pleistocene of Italy
(Calabria), the present species is more triangular in side view,
higher frontally, not truncated posteriorly, and punctate rather
than reticulated laterally. In contrast to Nipponocythere cola-
longoae (Ciampo, 1986), from the Upper Miocene of Italy
(Piedmont), this species is less elongated, not truncated poste-
riorly and less distincly rimmed and rather smooth along the
anterior margin. Our specimens also seem to be very close to
those referred to Cytheromorpha bimarginata Brestenská,
1975, described from the Lower Miocene (Egerian) of Slova-
kia and Hungary. The Middle Miocene material, however, is
more coarsely and more regularly punctate. It lacks a posterior
rib, bordering posteriorly lateral inflation, distally bounded by
less distinct parallel ribs, more pronounced in the LV. More-
over, our specimens have a much less pronounced posterodor-
sal rib, extending along the dorsal margin. Internal features,
including the hinge margin of Cytheromorpha bimarginata,
appear typical for Nipponocythere.
The material described by Říha (1983) from the Lower
Badenian of the Czech Republic, referred to Bosquetina car-
inella (Reuss, 1850), seems markedly different from those
representing B. carinella and falls within the variability of
Nipponocythere karsyensis sp. nov.
D i s t r i b u t i o n ( O c c u r r e n c e ) : Lower Badenian of the
Działoszyce Trough (Posądza) and the Korytnica Bay. Out-
side Poland – Lower Badenian of the Czech Republic (near
Brno) (Central Paratethys).
18 SZCZECHURA and AIELLO
Two ostracod species belonging to the genus Nippono-
cythere Ishizaki, 1971, have been found in the Middle Mi-
ocene (Badenian) deposits of the Polish part of the Fore-Car-
pathian Depression. Nipponocythere karsyensis sp. nov.
occurs in the Lower Badenian, while Nipponocythere silesien-
sis sp. nov. is characteristic of the Upper Badenian. In the
studied area, that is the northern part of the Central Paratethys,
Nipponocythere could have invaded in the early Middle Mi-
ocene (Lower Badenian) from the more southern parts of Eu-
rope, where they could be a remnant fauna of the former
Tethys, or they could have arrived, in the early Middle Mi-
ocene, from the Atlantic Ocean via a southern European route.
The Upper Badenian Nipponocythere silesiensis sp. nov. is
described from Upper Silesia, where it is associated with an
exceptionally abundant and diversified microfauna (various
groups, including ostracods). Apart from the ostracods, typi-
cal of the neritic and phytal (?) environment, there are numer-
ous “exotic” forms, which are extremely rare or (mostly) un-
known from the Middle Miocene outside the studied sections.
They are characterized by their small-size and thin-walled
shells. These specific features of this group of ostracods are
considered to be related to environmental conditions. The en-
vironment is considered to have been exceptionally favour-
able for ostracods, that is highly eutrophic, mostly with high
primary productivity of the surface waters. Such environmen-
tal conditions could have led to undersaturation in O
of the bottom waters, resulting in poor biocalcification
Acknowledgements: The authors are deeply thankful to Prof.
Dr. S.W. Alexandrowicz (Academy of Mining and Metallur-
gy, Kraków) for providing samples from the Gliwice bore-
holes: G-19 and G-21; to Dr. David J. Horne (School of Earth
and Environmental Sciences, University of Greenwich) for his
most valuable critical reading of the text as well as improving
the English language; to Dr. Wojciech Majewski (Institute of
Paleobiology of the Polish Academy of Sciences, Warszawa)
for his constructive remarks. Thanks also go to the reviewers
Dr. Radovan Pipík, Dr. Cemal Tunoglu, and especially to
Prof. Robin C. Whatley for his useful comments and improve-
ment of the manuscript. The SEM photomicrographs were
taken by Dr. J. Błaszyk (Institute of Paleobiology of the Pol-
ish Academy of Sciences, Warszawa), whereas the text figure
was prepared by A. Hołda, M.Sc, and plates by Ms. Danuta
Kościelska (the same Institute).
Aiello G. & Szczechura J. 2001a: First finding of the genus Nunana
McKenzie, Reyment & Reyment, 1993 (Paracytheridae: Ostra-
coda) from the Middle Miocene of Central Paratethys. Rev.
Esp. Micropaleont. 33, 1, 71—78.
Aiello G. & Szczechura J. 2001b: An ostracod marker for the Upper
Badenian (Miocene) of the Central Paratethys. Boll. Soc. Pale-
ont. Ital. 40, 1, 75—79.
Aiello G. & Szczechura J. 2002: Appearance of the genus Elofsonia
Wagner, 1957 (Loxoconchidae: Ostracoda) in the Middle Mi-
ocene of Central Paratethys: Rev. Micropaléont. 45, 1, 3—7.
Alexandrowicz S.W. 1963: Stratigraphy of the Miocene deposits in
the Upper Silesian Basin. Prace Inst. Geol. 39, 1—147 (in Pol-
ish with English summary).
Alexandrowicz S.W. 1997: Lithostratigraphy of the Miocene depos-
its in the Gliwice area (Upper Silesia, Poland). Bull. Pol. Acad.
Sci., Earth Sci. 45, 2—4, 167—179.
Ayress M. & Drapala V. 1996: New recent and fossil discoveries of
Cluthia (Ostracoda, Leptocytheridae) in the Southwest Pacific:
Implications on its origin and dispersal. In: Keen M.C. (Ed.):
Proc. 2nd Europ. Ostracod. Meet., Univ. Glasgow, Scotland
23rd-27th July, 1993. Brit. Microp. Soc., London, 149—158.
Babinot J.F., Carbonel P., Colin J.P., Peypouquet J.P. & Tambareau
Y. 1991: Variations morphologiques et adaptations morpho-
fonctionnelles chez les ostracodes: signification paleo-
environnementale. Geobios, Mem. Spec. 13, 135—145.
Baird W. 1850: The Natural History of the British Entomostraca.
Ray Soc., London, 1—364.
Bałuk W. & Radwański A. 1977: Organic communities and facies de-
velopment of the Korytnica Basin (Middle Miocene; Holy Cross
Mountains, Central Poland). Acta Geol. Pol. 27, 2, 85—123.
Barra D. 1995: First record of Nipponocythere Ishizaki, 1971 and
Swainocythere Ishizaki, 1981 (Ostracoda) in the Late Quater-
nary in the Mediterranean Basin. Boll. Soc. Paleont. Ital. 34, 2,
Barwicz-Piskorz W. 1997: Badenian (Miocene) Radiolaria from the
Gliwice area (Upper Silesia, Poland). Bull. Pol. Acad. Sci.,
Earth Sci. 45, 2—4, 87—95.
Benson R.H. 1976: Miocene deep-sea ostracodes of the Iberian Por-
tal and Balearic Basin. Mar. Micropal. 1, 3, 249—262.
Benson R.H. 1990: Ostracoda and the discovery of global Cainozoic
palaeoceanographical events. In: Whatley R. & Maybury C.
(Eds.): Ostracoda and Global Events. Brit. Micropal. Soc.
Publ. Ser., Chapman and Hall, London-New York-Tokyo-
Melbourne Madras, 41—58.
Bold W.A. van den 1985: Heinia, a new genus of Ostracoda from
the Gulf of Mexico and the Caribbean. J. Palaeont. 59, 1, 1—7.
Bonaduce G., Abate S. & Barra D. 1994: Kunihirella, a new marine
ostracod genus (Podocopa, Cytheruridae) from the Early
Pliocene of Eraclea Minoa (Sicily). Boll. Soc. Paleont. Ital. 33,
Braccini E. & Peypouquet J.P. 1996: Estimation of the intensity of
the oxygen minimum zone in the Santonian/Maastrichtian of
the southern margin of Tethys based on the analysis of archi-
tectural variability of ostracod shells. In: Keen M.C. (Ed.):
Proc. 2nd Europ. Ostracod. Meet., Univ. Glasgow, Scotland
23rd—27th July, 1993. Brit. Micropal. Soc., London, 67—74.
Brasier M.D. 1995: Fossil indicators of nutrient levels. 2: Evolution
and extinction in relation to oligotrophy. In: Bosence W.J. Dan
& Allison P.A. (Eds.): Marine Palaeoenvironmental Analysis
from Fossils. Geol. Soc. Spec. Publ. 83, 133—150.
Brestenská E. 1975: Ostracoden des Egerien. In: Brestenská E.
(Ed.): Chronostr. Neostrat., Miozän Zentralen Paratethys, 5
(OM: Egerien). VEDA, Bratislava, 377—435.
Brouwers E., Cronin T.M., Horne D.J. & Lord A.R. 2000: Recent
shallow marine ostracods from high latitudes: implications for
late Pliocene and Quaternary palaeoclimatology. Boreas 29,
Carbonel P. & Hoibian T. 1988: The impact of organic matter on os-
tracods from an equatorial deltaic area, the Mahakam delta –
southeastern Kalimantan. In: Hanai T., Ikeya N. & Ishizaki K.
(Eds.): Evolutionary biology of Ostracoda – its fundamentals
and applications. Proc. 9th Intern. Symp. on Ostracoda, Shi-
zuoka, Japan, 29 July-2 August 1985, Developments in Palae-
ont. and Stratigr. 11, Elsevier, Amsterdam-Oxford-New
THE OSTRACOD GENUS NIPPONOCYTHERE ISHIZAKI, 1971 19
Carbonnel P. & Ballesio R. 1982: Les ostracodes Pliocènes du sud-
est de la France. Docum. Lab. Géol. Lyon. 85, 1—113.
Ciampo G. 1986: Ostracods fromTortonian/Messinian boundary in
some Italian sections. Boll. Soc. Paleont. Ital. 24, 1, 29—110
Colalongo M.L. & Pasini G. 1980: The Plio/Pleistocene ostracod
fauna of Vrica section in Calabria (with considerations on the
Neogene/Quaternary boundary). Boll. Soc. Paleont. Ital. 19, 1,
44—126 (in Italian).
Coles G.P., Ainsworth N.R., Whatley R. & Jones R.W. 1996: Fora-
minifera and Ostracoda from Quaternary carbonate mounds as-
sociated with gas seepage in the Porcupine Basin, offshore
western Ireland. Rev. Esp. Micropal. 28, 2, 113—151.
Coles G., Ayress M. & Whatley R. 1990: A comparison of North
Atlantic and Pacific Cainozoic deep-sea Ostracoda. In: What-
ley R. & Maybury C. (Eds.): Ostracoda and Global Events.
Brit. Micropal. Soc. Publ. Ser., Chapman and Hall, London—
New York—Tokyo—Melbourne—Madras, 287—305.
Cronin T.M., DeMartino D.M., Dwyer G.S. & Rodriguez-Lazaro J.
1999: Deep-sea ostracode species diversity: response to Late
Quaternary climate change. Marine Ostracoda and Global
Change. Mar. Micropal., Spec. Issue 37, 3—4, 231—249.
Curry B.B. 1999: An environmental tolerance index for ostracodes
as indicators of physical and chemical factors in aquatic habi-
tats. In: Holmes J.A. & Horne D.J. (Eds.): Non-marine Ostra-
coda: Evolution and environment. (Theme 1, 13th Intern.
Symp. on Ostracoda (ISO97)). Palaeogeogr. Palaeoclimat.
Palaeoecol., Spec. Issue 148, 1-3, 51—63.
Dingle R.V. 1995: Continental shelf upwelling and benthic Ostraco-
da in the Benuela System (southeastern Atlantic Ocean). Mar.
Geol. 122, 207—225.
Drapala V. & Ayress A. 1993: On Nipponocythere colalongoae (Ci-
ampo): In: Athersuch J., Horne D.J., Siveter D.J. & Whittaker
J.E. (Eds.): A Stereo-Atlas of Ostracod Shells. 20, 5, 17—24.
Durakiewicz T., Gonera M. & Peryt T.M. 1997: Oxygen and carbon
isotopic changes in the Middle Miocene (Badenian) Foramin-
ifera of the Gliwice area (SW Poland). Bull. Pol. Acad. Sci.,
Earth Sci. 45, 2—4,145—156.
Funnell B.M., Haslett S.K., Kennington K., Swallow J.E. & Kersley
C.L. 1996: Strangeness of the equatorial Ocean during the Old-
uvai magnetosubchron (1.95 to 1.79 Ma). In: Moguilevsky A.
& Whatley R. (Eds.): Microfossils and Oceanic Environments.
Proc. of the “ODP and the Marine Biosphere” Int. Confer-
ence, Aberystwyth, 19th-21st April 1994, 93—109.
Garlicki A. & Szybist A. 2001: Organic matter of Miocene sedi-
ments within the overburden of salt deposit Wojnicz near Tar-
now. Geologia 27, 2—4, 341—361 (in Polish with English
Gedl P. 1997: Palynofacies of the Miocene deposits in the Gliwice
area (Upper Silesia, Poland)). Bull. Pol. Acad. Sci., Earth Sci.
45, 2—4, 191—210.
Gonera M. 1997: Miocene foraminiferal assemblages in the Gliwice
area. Bull. Pol. Acad. Sci., Earth Sci. 45, 2—4, 97—105.
Gooday A.J. 1994: The biology of deep-sea foraminifera: A review
of some advances and their applications in paleoceanography.
Palaios 9, 1, 14—31.
Górka H. 1997: Holothurian sclerites from the Miocene deposits of
the Gliwice area (Upper Silesia, Poland). Bull. Pol. Acad. Sci.,
Earth Sci. 45, 2—4, 105—118.
Hanai T. 1957: Studies on the Ostracoda from Japan: 2. Subfamily
Pectocytherinae n. subfam. J. Fac. Sci. Univ. Tokyo. 2, 10,
Hutchings L., Pitcher G.C., Probyn T.A. & Bailey G.W. 1995: The
chemical and biological consequences of coastal upwelling. In:
Summerhayes C.P., Emeis K.-C., Angel M.V., Smith R.L. &
Zeitzschel B. (Eds.): Upwelling in the ocean: Modern process-
es and ancient records. Wiley & Sons, Chichester—New York—
Ishizaki K. 1971: Ostracodes from Aomori Bay, Aomori Prefecture,
Northeast Honshu, Japan. Tohoku Univ., Sci. Rep. 2nd Ser. 43,
Ishizaki K. 1981: Ostracods from the East China Sea. Tohoku Univ.,
Sci. Rep., 2nd Ser. 51, 1—2, 37—65.
Janssen A.W. & Zorn I. 1993: Revision of Middle Miocene hol-
oplanktonic gastropods from Poland, published by the late
Wilchelm Krach. Scripta Geol., Spec. Issue 2, 155—236.
Kaiho K. 1994: Planktonic and benthic foraminiferal extinction
events during the last 100 m.y. Palaeogeogr. Palaeoclimat.
Palaeoecol. 111, 4, 45—71.
Kaiho K. 1999: Effect of organic carbon flux and dissolved oxygen
on benthic foraminiferal oxygen index (BFOI). Mar. Micropal.
37, 1, 67—76.
Kowalewski M. & Miśniakiewicz W. 1993: Reliability of quantita-
tive data on fossil assemblages: a model, a simulation, and an
example. Neu. Jb. Geol. Paläont., Abh. 187, 243—260.
Łuczkowska E. 1974: Micropaleontological stratigraphy of Mi-
ocene in the light of the VIIth Symposium on the Paratethys
CMNS in Cracow. Sprawozd. pos. Komisji Nauk., Polska Aka-
demia Nauk, Warszawa-Krakow, 18, 2, 499—501 (in Polish).
Łuczkowska E. 1998: Marine Miocene deposits of the Paratethys in
Poland. In: Cícha I., Rögl F., Rupp Ch. & Čtyroká I. (Eds.):
Oligocene-Miocene foraminifera of the Central Paratethys.
Abh. Senckenberg. Naturforsch. Gesell. 549, 28—34.
Malz H. 1981: Nipponocythere (Ostracoda), eine inverse loxo-
conchine Gattung: Senckenberg. Lethaea 62, 2, 155—165.
Maddocks R.F. & Steineck P.L. 1987: Ostracoda from experimental
wood-island habitats in the deep sea. Micropaleontology 33, 4,
McKenzie K.G. 1974: Cenozoic Ostracoda of southeastern Austra-
lia with the description of Hanaiceratina new genus. Geo-
science and Man 6, 153—182.
McKenzie K.G., Reyment R.A. & Reyment E.R. 1993: Eocene Os-
tracoda from the Browns Creek Clays at Browns Creek and
Castle Cove, Victoria, Australia. Rev. Esp. Paleont. 8, 75—116.
Mezquita F., Tapia G. & Roca J.R. 1999: Ostracoda from springs on
the eastern Iberian Peninsula: ecology, biogeography and pale-
olimnological implications. In: Holmes J.A. & Horne D.J.
(Eds.): Non-marine Ostracoda: Evolution and environment.
(Theme 1, 13th Itern. Symp. on Ostracoda (ISO97)). Palaeo-
geogr. Palaeoclimat. Palaeoecol., Spec. Issue 148, 1-3, 65—85.
Murray J.W. 1991: Ecology and palaeoecology of benthic Foramin-
ifera. Longman, Harlow, 1—397.
Murray J.W. 1995: Microfossils indicators of ocean water masses,
circulation and climate. In: Bosence W.J. Dan & Allison P.A.
(Eds.): Marine palaeoenvironmental analysis from fossils. Geol.
Soc. Spec. Publ. 83, 245—264.
Nascimento A. & Říha J. 1996: Ostracod species common to the
Neogene of Portugal and Central Paratethys. In: Keen M.C.
(Ed.): Proc. 2nd Europ. Ostracod. Meet., Univ. Glasgow,
Scotland 23rd-27th July, 1993. Brit. Microp. Soc., Lon-
Neale J.W. 1971: Microfaunas and some aspects of the Speeton
Clay environment. In: Oertli H.J. (Ed.): Paleoecologie des Os-
tracodes. Colloque Pau (1970). Bull. Centre Res. Pau-SNPA,
Suppl. 5, 663—681.
Ney R., Burzewski W., Bachleda T., Górecki W., Jakóbczak K. &
Słupczyński K. 1974: Outline of paleogeography and evolution
of lithology and facies of Miocene layers on the Carpathian
Foredeep. Prace Geol. PAN 82, 7—64 (in Polish with English
Oertli H.J. 1971: The aspect of ostracode faunas – A possible new
tool in petroleum sedimentology. In: Oertli H.J. (Ed.): Paleo-
20 SZCZECHURA and AIELLO
ecologie des Ostracodes. Colloque Pau (1970). Bull. Centre
Res. Pau-SNPA, Suppl. 5, 137—150.
Osmólski T. 1972: The influence of the geological structure of the
marginal parts of the Działoszyce Trough on the metasomato-
sis of gypsum. Biul. Inst Geol. 260, 65—188 (in Polish with En-
Peryt D. 1997: Calcareous nannoplankton stratigraphy of the Mid-
dle Miocene in the Gliwice area (Upper Silesia, Poland). Bull.
Pol. Acad. Sci., Earth Sci. 45, 2—4, 119—131.
Peypouquet J.-P. 1977: Les Ostracodes et la connaissance des pale-
omilieux profonds. Application au Cenozoic de l’Atlantique
nord-oriental. These Doctor. Etat Sci., Universite Bordeaux I,
Peypouquet J.P., Carbonel P., Ducasse O., Tolderer-Farmer M. &
Lete C. 1988: Environmentally cued polymorphism of ostracods.
In: Hanai T., Ikeya N. & Ishizaki K. (Eds.): Evolutionary biolo-
gy of Ostracoda – its fundamentals and applications. Proc. 9th
Intern. Symp. on Ostracoda, Shizuoka, Japan, 29 July-2 August
1985. Developments in Palaeont. and Stratigr. 11, Elsevier, Am-
sterdam—Oxford—New York—Tokyo, 1003—1019.
Radwańska U. 1992: Fish otoliths in the Middle Miocene (Bade-
nian) deposits of southern Poland. Acta Geol. Pol. 42, 3—4,
Reuss A.M. 1850: Die Fossilen Entomostraceen des osterreichis-
chen Tertiarbeckens. Naturwiss. Abh. 3, 1, 41—92.
Říha J. 1983: Ostrakoden aus der kalkigen tone (tegels) des unteren
Badenien (Tertiär-Miozän) in umgebung der stadt Brünn. Acta
Mus. Morav., Sci. Natur. 68, 57—83 (in Czech).
Říha J. 1989: Ostracod interpretation of palaeodepth of Miocene
(Lower Badenian) calcareous clays near Brno, Czechoslova-
kia. Cour. Forsch.-Inst. Senckenberg. 113, 103—116.
Rodriguez-Lazaro J. & Garcia Zarraga E. 1996: Paleogene deep-
marine ostracodes from the Basque Basin. In: Keen M.C. (Ed.):
Proc. 2nd Europ. Ostracod. Meet., Univ. Glasgow, Scotland
23rd-27th July, 1993. Brit. Micropal. Soc., London, 79—85.
Rögl F. 1998: Palaeogeographic considerations for Mediterranean
and Paratethys seaways (Oligocene to Miocene). Ann.
Naturhist. Mus. Wien 99A, 279—310.
Rögl F. & Brandstötter F. 1993: The foraminifera genus Amphiste-
gina in the Korytnica Clays (Holy Cross Mts, Central Poland)
and its significance in the Miocene of the Paratethys. Acta.
Geol. Pol. 43, 1—2, 97—108.
Sars G.O. 1866: Oversigt af Norges marine ostracoder. Förhandl.
Vidensk. Selskab Christiania 7, 1—130.
Sars G.O. 1925 (1922—1928): An account on the Crustacea of Nor-
way. Bergen Museum (Norway) 9, 1—277.
Sen Gupta B.K. & Machain-Castillo M.L. 1993: Benthic foramin-
ifera in oxygen-poor habitats. Mar. Micropal. 20, 2, 183—201.
Steineck P.L, Maddocks R.F., Coles G. & Whatley R. 1990: Xylo-
phile Ostracoda in the deep sea. In: Whatley R. & Maybury
C. (Eds.): Ostracoda and global events. Brit. Micropal. Soc.
Publ. Ser., Chapman & Hall, London—New York—Tokyo—
Studencka B., Gontsharova I.A. & Popov S.Y. 1998: The bivalve
faunas as a basis for reconstruction of the Middle Miocene his-
tory of the Paratethys. Acta Geol. Pol. 48, 3, 282—342.
Summerhayes C.P., Emeis K.-C., Angel M.V. & Smith R.L. 1995:
Upwelling in the ocean: Modern processes and ancient records.
In: Summerhayes C.P., Emeis K.-C., Angel M.V., Smith R.L.
& Zeitzschel B. (Eds.): Upwelling in the ocean: Modern pro-
cesses and ancient records. Wiley & Sons, Chichester—
New York— Brisbane—Toronto—Singapore, 1—38.
Szczechura J. 1982: Middle Miocene foraminiferal biochronology
and ecology of SE Poland. Acta Pal. Pol. 27, 1—4, 3—44.
Szczechura J. 1986: Cluthia miocenica sp. n. from the Middle Mi-
ocene of southern Poland (Central Paratethys). J. Micropal. 5,
Szczechura J. 1994: Paleogeographic meaning of psychrospheric
Miocene ostracodes from the Fore-Carpathian Depression.
Acta Geol. Pol. 44, 1—2, 138—151.
Szczechura J. 1995: The ostracod genus Xylocythere Maddocks &
Steineck, 1987, from the Middle Miocene of the Fore-Car-
pathian Depression, southern Poland (Central Paratethys),
and its biogeograhic significance. Acta Geol. Pol. 45, 1—2,
Szczechura J. 1996: Ecological control of ostracode distribution in
the Middle Miocene of Paratethys. In: Keen M.C. (Ed.): Proc.
2nd Europ. Ostracod. Meet. Univ. Glasgow, Scotland, 23rd-
27th July, 1993. Brit. Micropal. Soc., London, 99—106.
Szczechura J. 1997: Bolboforms (Protophyta, incertae sedis) from
the Middle Miocene of Upper Silesia (Carpathian Foredeep).
Bull. Pol. Acad. Sci., Earth Sci. 45, 2—4, 133—144.
Szczechura J. 1998: The distribution of Triebelina raripila and Car-
inocythereis carinata (Ostracoda) from the Middle Miocene of
the Central Paratethys and their paleogeographic implications.
J. Micropal. 17, 2, 125—130.
Szczechura J. 2000: Palaeoenvironments of the Middle Miocene
evaporite-bearing deposits from the Działoszyce Trough, Car-
pathian Foredeep, Poland, based on microfaunal studies. Geol.
Quart. 44, 2, 119—135.
Van Harten D. 1990: Modern abyssal ostracod faunas of the eastern
Mid-Atlantic Ridge area in the North Atlantic and a compari-
son with the Mediterranean. In: Whatley R. & Maybury C.
(Eds.): Ostracoda and global events. Brit. Microp. Soc. Publ.
Ser., Chapman and Hall, London—New York—Tokyo—Mel-
Van Harten D. 1992: Hydrothermal vent ostracoda and faunal asso-
ciation in the deep sea. Deep Sea Res. 37, 6, 1067—1070.
Van Harten D. 1993: Deep sea hydrothermal vent eucytherurinae
Ostracoda: the enigma of the pore clusters and paradox of the
hinge. In: McKenzie K.G. & Jones P.J. (Eds.): Ostracoda in the
Earth and Life Sciences. Proc. 11th Intern. Symp. Ostracoda,
Warnambool, Victoria, Australia July/August 1991, Balkema,
Whatley R.C. & Ayress M. 1988: Pandemic and endemic distribu-
tion patterns in Quaternary deep-sea Ostracoda. In: Hanai T.,
Ikeya N. & Ishizaki K. (Eds.): Evolutionary biology of Ostra-
coda – its fundamentals and applications. Proc. 9th Intern.
Symp. on Ostracoda, Shizuoka, Japan, 29 July-2 August 1985.
Developments in Palaeont. and Stratigr. 11, Elsevier, Amster-
dam—Oxford—New York—Tokyo, 739—755.
Whatley R., Eynon M. & Moguilevsky A. 1996: Recent Ostracoda
of the Scoresby Sund fjiord system, East Greenland. Rev. Esp.
Micropal. 28, 2, 5—23.
Whatley R., Staunton M. & Kaesler R.L. 1997: The depth distribu-
tion of recent marine Ostracoda from the southern Strait of Ma-
gellan. J. Micropal. 16, 121—130.
Witkowski A. & Gonera M. l997: Miocene diatom flora from Cen-
tral Paratethys (Carpathian Foredeep, Upper Silesia) – prelim-
inary results. Bull. Pol. Acad. Sci., Earth Sci. 45, 2—4, 181—190.