GeolCarp_Vol54_No1_9

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

JANINA SZCZECHURA

and GIUSEPPE AIELLO

Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warszawa, Poland; janina.s@twarda.pan.pl

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

and/or O

Key words: Middle Miocene, Fore-Carpathian Depression, paleoecology, paleogeography, Ostracoda.

Introduction

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
(abbreviated ZPAL).

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
foraminifers.

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
indices.

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-
posit.

Gliwice G-19 and G-21 boreholes (Działoszyce

Trough)

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
less abundant.

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-
wellings.

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

Paleozoogeography

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-
sentatives

of Heinia

(another

junior

synonym

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
Hemisphere.

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-
tral Paratethys.

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
Atlantic.

Paleoecology

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
climate.

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
typical habitat.

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
Fore-Carpathian Depression.

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
shells.

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
chemical parameters.

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
CaCO

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-

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.

Systematic descriptions

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-

dle Miocene).

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

Silesia.

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
margin.

M a t e r i a l : 15 adult valves and one adult carapace, well

preserved.

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
dorsal margin.

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 ) :

ZPAL O.48/60

holotype

ZPAL O.48/62

paratype

ZPAL O.48/61

paratype

RV LV LV

Length 0.44

Length 0.39

Height 0.23

Height 0.20

R e m a r k s : This species is similar to that described by

Drapala & Ayress (1993), from the Late Quaternary of Aus-

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
branch anteriorly.

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.

(Fig. 2.1—5)

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-

dle Miocene).

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

Karsy.

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

preserved.

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 ) :

ZPAL O.48/319,

holotype

ZPAL O.48/305,

paratype

ZPAL O.48/318,

paratype

LV LV RV

Length 0.38

Length 0.36

Length 0.40

Height 0.23

Height 0.25

Height 0.23

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

Conclusions

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

and/or

CaCO

of the bottom waters, resulting in poor biocalcification

of ostracods.

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).

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