www.geologicacarpathica.sk
GEOLOGICA CARPATHICA, OCTOBER 2009, 60, 5, 419—430 doi: 10.2478/v10096-009-0030-3
Introduction
The Mediterranean Sea and the Paratethys were formed as
new marine realms during the Late Eocene (Popov et al.
1993; Rögl 1999). These basins underwent complex paleo-
geogeographical changes especially in the Miocene, such as
the opening and closing of marine connections with the Indi-
an Ocean to the East and the Mediterranean Sea and the At-
lantic Ocean to the West (Rögl 1999, 2001).
Strong geographical separation also took place within the
Paratethys. Basin formation led to semi-independent evolu-
tion of the Western (from the Rhone Basin to Western Ba-
varia), Central (from Bavaria to the eastern foredeep of the
Carpathian) and Eastern (from the Euxinian basin complex
to Lake Aral) Paratethyan realms (Rögl 1999; Steininger &
Wessely 2000; Piller & Harzhauser 2005).
In particular, in the Middle Miocene a regressive phase took
place at the end of the Badenian (Central Paratethys) corre-
sponding to the Konkian (Eastern Paratethys) leading to a dra-
matic change in marine biota (Harzhauser & Piller 2007). A
restriction of the connections between the Paratethys and the
open ocean occurred at the beginning of the latest Middle Mi-
ocene and the connections with the Indo-Pacific disappeared
(Rögl 1999; Harzhauser & Piller 2007). In the Middle—Late
Miocene the Paratethys was composed of different semi-en-
closed sub-basins including the Pannonian in the Central
Paratethys and Euxinic—Caspian Basins in the Eastern Para-
tethys (Kolesnikov 1935). Thus, the complex evolution of the
Paratethys is reflected in the distribution and evolution of the
fauna and flora with the proliferation of foraminifera, differing
in abundance and composition from those of open oceans (e.g.
Iaccarino 1985; Rögl 1985; Cicha et al. 1998).
This setting makes correlation of sediments between these
realms both interesting and problematic. In the past, different
Taxonomic revision and new species/subspecies of
Middle-Late Miocene (Bessarabian) miliolids of the Family
Hauerinidae Schwager from Georgia – Eastern Paratethys
LAMARA MAISURADZE
1
, KAKHABER KOIAVA
2
and SILVIA SPEZZAFERRI
3
1
The Georgian National Museum, L. Davitashvili Institute of Paleobiology, Tbilisi, Georgia; lamaramaisuradze@yahoo.com
2
The Georgian Academy of Science, Alexandre Djanelidze Institute of Geology, Tbilisi, Georgia; koiava_ka@yahoo.com
3
University of Fribourg, Department of Geosciences, Ch. du Musée 6, 1700 Fribourg, Switzerland; silvia.spezzaferri@unifr.ch
(Revised manuscript received March 16, 2009; accepted in April 20, 2009)
Abstract: Three new miliolid taxa from Bessarabian sediments from Georgia (Eastern Paratethys) are described fol-
lowing the classification of Łuczkowska (1972), which has never been used before by ex-Soviet micropaleontologists.
They are: Varidentella luczkowskae; Varidentella reussi (Bogdanowicz) subsp. costulata; and Affinetrina voloshinovae
(Bogdanowicz) subsp. eldarica. This classification takes into account the morphology of the aperture and the shape and
size of teeth as criteria to distinguish the species. This study contributes important criteria which will help to unify the
taxonomical inconsistencies between the Eastern, Central and Western Paratethyan miliolids.
Key words: Bessarabian, Georgia, Paratethys, Foraminifera, miliolid.
chrono/geochronological time scales were applied in studies
of these two basins (e.g. Steininger & Wessely 2000). The
lack of dialogue between western and eastern countries, has
led to two independent sets of geological and stratigraphical
correlations. In particular, the Sarmatian regional stage is de-
fined differently in the Western and Central with respect to the
Eastern Paratethys (e.g. Steininger & Wessely 2000; Piller &
Harzhauser 2005). Figure 1 shows the stratigraphic correlation
between Middle-Late Miocene Mediterranean, Central and
Eastern Paratethys regions.
Fig. 1. Stratigraphic correlation between Mediterranean and Para-
tethyan stages. The Sarmatian regional stage spans a longer time in
the eastern basin. Modified after Piller & Harzhauser (2005) and
Harzhauser & Piller (2007). The regional stages Volhynian = N
1
S
1
;
Bessarabian = N
1
S
2
; Khersonian = N
1
S
3
after Koiava (2006).
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MAISURADZE, KOIAVA and SPEZZAFERRI
Previous studies on the Family Hauerinidae
Miliolids belonging to the Family Hauerinidae are very
abundant in Miocene deposits from the Paratethys and are use-
ful for dating and characterizing these sediments. In particular,
their consistent regional development and their distribution,
seem to be related to the local/regional paleoenvironmental
conditions of the different sub-basins of the Paratethyan
realms (Maisuradze 1971).
The classification of the Family Hauerinidae Schwager
(1876) is very complex and is still being debated. D’Orbigny
(1826) separated this group of miliolids into five genera
Biloculina, Triloculina, Quinqueloculina, Spiroloculina and
Adelosina based on the number and shape of the chambers in
the last whorl. Williamson (1858) grouped the genera Quin-
queloculina d’Orbigny and Triloculina d’Orbigny into one
single genus Miliolina. Brady (1884) also included Adelosi-
na d’Orbigny into the genus Miliolina Williamson. Schlum-
berger (1887, 1893) described the two genera Sigmoilina and
Massilina. Wiesner (1931) included them in the genus Milio-
lina Williamson. However, he separated the genus Milio-
linella characterized by a flat, plate-like tooth. Bogdanowicz
(1947, 1952) followed Brady’s (1884) classification, but also
included Massilina Schlumberger in the genus Miliolina
Williamson. In addition Bogdanowicz (1960) proposed, that
the amplitude of the angle between the successive chambers
should be the main criteria for diving miliolids and restored
the genera Quinqueloculina d’Orbigny and Triloculina
d’Orbigny, characterized by an angle of 72°, and 120° be-
tween successive chambers, respectively. Vella (1957) di-
vided the genus Quinqueloculina d’Orbigny into three
subgenera: Quinqueloculina (Quinqueloculina) d’Orbigny,
Quinqueloculina (Lachlanella) Vella, Quinqueloculina sen-
su lato which groups the remaining forms based on the wall
texture, tooth size and shape of aperture.
Since those pioneering studies the taxonomy of miliolids
has been revised many times (e.g. Loeblich & Tappan 1964),
and the currently accepted taxonomy was proposed by Loe-
blich & Tappan (1988). However, Ex-USSR micropaleontol-
ogists have always used the “old” criteria suggested by
Bogdanowicz (1959, 1981) for taxonomic division, since
western literature is not available in their region.
Łuczkowska (1972, 1974) extensively studied the miliolid
fauna from Miocene deposits of Poland, and, at the beginning
of her studies, applied the classification of Loeblich & Tappan
(1964). However, in the course of her research, she became
aware of some inconsistencies concerning the main diagnostic
criteria. For example, the main feature to separate the various
genera in the Subfamilies Quinqueloculininae Cushman, Mil-
iolinae Ehrenberg and Miliolinellininae Vella is the outline of
the aperture, but in the Subfamilies Fabulariinae Reuss and
Tubinellinae Rhumbler it is the internal construction of the
tests. As a result, species that were characterized by similar
morphological features but had different shaped teeth were in-
cluded in the Subfamilies Quinqueloculininae Cushman, Mil-
iolinae Ehrenberg and Miliolinellininae Vella and were
considered to be phylogenetically related. Furthermore, gen-
era with a non-quinqueloculiniform coiling mode such as
Flintina Cushman, Sigmoilina Schlumberger, Ptychomiliolla
Eimer & Fickert, Welmanella Finlay were included in the
Subfamily Quinqueloculininae Cushman. In the Subfamilies
Miliolinellinae Vella and Miliolinae Ehrenberg forms were in-
cluded, which were coiled like typical quinqueloculinids.
Therefore, Łuczkowska (1972, 1974) proposed a new clas-
sification of miliolids based on the inner structure of the
shell, the general morphology, the shape and size of the ap-
erture and the tooth. She identified four types of aperture
typical for Miocene miliolids. (1) Wide, oval-elongated,
with a simple-shaped long tooth, sometimes bifid at the end;
(2) high, narrow, with parallel edges and an elongated tooth;
(3) Closed, at the base of the last chamber, with a short V-like
tooth; (4) Open semicircular, placed at the base of the last
chamber and characterized by a quadrangular tooth, which
sometimes may be reduced.
She also identified two subfamilies that include genera
still living today, based on the inner structure of the test, the
outline of the aperture and the shape of the tooth. (1) Sub-
family Quinqueloculininae Cushman emended Łuczkowska
(1972), this group includes quinqueloculiniform species with
all the typical characters of the genus Quinqueloculina.
These forms possess, a 2-layer wall texture, an oval and
elongate aperture, a long simple or bifid tooth, and an angle
of 140°
between the successive chambers. Five and three
chambers can be observed on the two sides of the test respec-
tively. Beside the genus Quinqueloculina sensu stricto this
subfamily, included the two genera Lachlanella proposed by
Vella (1957) characterized by a narrow, elongated and fis-
sure-like aperture with parallel lips and a simple elongated
tooth; and Cycloforina Łuczkowska characterized by a circu-
lar aperture and a short tooth. (2) Subfamily Miliolinellinae
Vella (1957) emended Łuczkowska (1972). This second
group comprises forms with a crypto-quinqueloculinid coil-
ing mode. In these forms only three chambers are visible in
the external whorl, however two smaller and concealed
chambers are present which complete the quinqueloculinid
coiling mode. Adult forms of this group display a triloculin-
id coiling mode, in which the chambers of the last whorl are
placed at 120°
from each other. In this group, forms are also
included which have rounded chambers coiled in planes at
130° (or more than 130°)
and the last chamber coiled at 90°
with respect to the previous chambers.
These forms were grouped into four new genera: (1) Vari-
dentella Łuczkowska with a semicircular oblique aperture
characterized by a low and flat tooth; (2) Affinetrina Łucz-
kowska with a longitudinal slit-like aperture and a long tooth
similar to that of Lachlanella Vella; (3) Sinuloculina Łucz-
kowska with a circular aperture, a short bifurcate tooth, simi-
lar to that in Triloculina d’Orbigny and Cycloforina
Łuczkowska, and the chambers of the last whorl coiled at
180°. (4) Crenatella Łuczkowska, has a narrow slit-like ap-
erture with serrate edges, a crescent-like, oblique tooth in the
aperture and additionally the inner structure of the test is
similar to the structure observed in the Family Miliolina
Ehrenberg.
The classification of Łuczkowska (1972, 1974) presents
some limitations. In particular, for its correct application it is
often necessary to study thin sections that show the inner
structure of the test, otherwise identification of high taxo-
421
HAUERINIDAE SCHWAGER – TAXONOMIC REVISION OF BESSARABIAN MILIOLIDS (EASTERN PARATETHYS)
nomic units like subfamilies and families is not possible.
Thin sections are not always available. Additional problems
arise because, in genera such as Varidentella Łuczkowska,
Hauerina d’Orbigny and Miliola Lamarck, changes of both
size and shape of the aperture as well as the tooth can occur
at different stages of the ontogeny. Ontogenetic changes im-
ply that transitional forms must be classified according to
their external morphology (e.g. shape of aperture, teeth and
number of chambers). Apart from these limitations the clas-
sification of Łuczkowska (1972, 1974) remains the most
complete and reliable framework to describe and classify
Paratethyan miliolids.
Loeblich & Tappan (1988) accepted most of the genera
described by Łuczkowska (1972, 1974). However, in their
Fig. 2. Geological map of the areas of Ckhorocku and Martkophi, where the Ochkhamuri and Satskhenisi sections outcrop.
422
MAISURADZE, KOIAVA and SPEZZAFERRI
classification they placed many of them into the Family
Hauerinidae Schwager. According to Loeblich & Tappan
(1988) this family includes forms with a proloculus followed
by two chambers, rarely with intervening flexostyle. Cham-
bers are added in one to five or more planes of coiling. Rare-
ly the adult test may have more than two chambers per whorl
or may be uncoiled and rectilinear. The aperture placed at
the end of the last chamber may have a simple or a complex
tooth. This family differs from the Family Miliolidae Ehren-
berg because the genera included in this latter present nu-
merous perforations or pseudopores in the wall.
Presented here are the descriptions of one new species and
two sub-species according to the classification of Łuczkowska
(1972, 1974) modified by Loeblich & Tappan (1988). The ap-
plication of this classification, allows a better comparison be-
tween Western, Central and Eastern Paratethyan species and
contributes to unifying the taxonomy between the two regions.
Materials and methods
The benthic foraminiferal assemblages analysed in this
study come from two sections in Western and Eastern Geor-
gia, respectively (Koiava 2006). The Ochkhamuri section is
located in the gorge of river Ochkhamuri in Western Georgia,
at a distance of 1 km from the village of Lesichino, in the
gorge of the river Ochkhamuri. Sediments span the regional
stage N
1
S
2
(Koiava 2006) = Bessarabian. The Satskhenisi
section is located in the gorge of the river Satskhenisi about
1—1.5 km north-west from the village of Satskhenisi (Eastern
Georgia). Sediments span the regional stage N
1
S
1
= Vol-
hynian, and N
1
S
2
(Koiava 2006) = Bessarabian (Fig. 2).
Samples were washed using standard techniques for fora-
miniferal preparation (e.g. Spezzaferri & Ćorić 2001). The
entire collection is presently stored at the Georgian National
Museum, L. Davitashvili Institute of Paleobiology or at the
A. Djanelidze Institute of Geology, in Tbilisi. Scanning
Electron Microscope (SEM) images were obtained with the
FEI XL30 Sirion FEG microscope at the University of Fri-
bourg, Department of Geosciences.
Sedimentology
The lithology and the studied samples together with the thick-
ness of the five distinct layers that can be observed in the Och-
khamuri and Satskhenisi sections are reported in Figs. 3 and 4.
The Ochkhamuri section consists of 5 layers: Layer 1 is 9 m
thick and consists of bluish-grey clays with sandy clay layers;
Layer 2 is 3 m thick and consists of bluish-grey clays with in-
tercalation of sandstone layers; Layer 3 is 5 m thick and is
characterized by alternating clay and sandstone layers; Lay-
er 4 is 5 m thick and consists of bluish-grey sandy clays;
Layer 5 is 6 m thick and consists of grey marls.
The Satskhenisi sections consist of 6 layers: Layer 1 is
80 m thick and consists of bluish-grey clays with intercala-
tions of yellowish-grey sandstones and greyish sandy clays;
Layer 2 is 57 m thick and consists of bluish-grey clays with
interlayered yellowish-grey sandstones and greyish sandy
clays at the lower parts of it; Layer 3 is 7 m thick and con-
sists of bluish-grey clays; Layer 4 is 20 m thick and consists
of bluish-grey clays; Layer 5 is 3 m thick and consists of yel-
lowish-grey sandstones; Layer 6 is 95 m, it consists of blu-
ish-grey clays with interbedded yellowish-grey sandstones.
Taxonomy
Maisuradze (1971) informally described a few species of
Bessarabian miliolids from Georgia but left them in the open
nomenclature. These species are now officially described ac-
cording to Łuczkowska (1972, 1974) with modification by
Loeblich & Tappan (1988) and presented for the first time in
western literature. Their distribution and accompanying as-
semblages are documented in Figs. 3 and 4.
Suborder: Miliolina Delage & Hérouad, 1896
Superfamily: Miliolacea Ehrenberg, 1839
Family: Hauerinidae Schwager, 1876
Subfamily: Hauerininae Schwager, 1876
Genus: Varidentella Łuczkowska, 1972
Varidentella reussi costulata subsp. nov.
Fig. 5.1a—c,2a—c,3a—c
1971 Quinqueloculina sp. Maisuradze, pl. IV, figs. 3a—c; 4a—c
no 1974 Varidentella georgiana Łuczkowska, p. 138, pl. XXVII,
figs. 3—4
no 1998 Varidentella reussi Cicha et al., p. 136, pl. 17, figs. 13—14
M o t i v a t i o n : Maisuradze (1971) described this form as
belonging to the Q. reussi group. However, the main differ-
ence between typical V. reussi and the taxon described here
is the presence of numerous striae as ornamentation of the
wall texture.
In particular, Maisuradze (1971) identified this subspecies
and termed it Quinqueloculina sp. and Łuczkowska (1974)
described it as Varidentella georgiana. However, V. georgi-
ana differs from the forms described in Maisuradze (1971)
by the general outline of the test. Varidentella georgiana is
rounded, the chambers are strongly arched forming semicir-
cles and the proximal part near the aperture is strongly con-
vex. Additionally, the tooth is slightly oval. The outline of
Quinqueloculina sp. as described by Maisuradze (1971) is
wide-oval, the tooth is wide and sub-quadrangular more sim-
ilar to the V. reussi group. Therefore, V. georgiana and
Quinqueloculina sp. of Maisuradze (1971) should be consid-
ered as two distinct taxa and Quinqueloculina sp. should be
regarded as related to V. reussi because of its strong resem-
blance to this species. Additionally, V. georgiana Łuczkows-
ka is a typical form from the Lower Sarmatian (Volhynian)
deposits of the Western and Central Paratethys. According to
our data, V. reussi costulata first occurs only in Bessarabian
deposits of Georgia, Crimea, Moldova and Pre-Caucasus.
H o l o t y p e : It is presented in Fig. 5.1a—c, and it is stored
at the Georgian National Museum, L. Davitashvili Institute
of Paleobiology with the reference number 14/M.
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HAUERINIDAE SCHWAGER – TAXONOMIC REVISION OF BESSARABIAN MILIOLIDS (EASTERN PARATETHYS)
Fig. 3.
Lithological
log
and
distribution
of
benthic
foraminifers
in
the
Ochkhamuri
section.
N
1
S
2
=
Bessarabian.
424
MAISURADZE, KOIAVA and SPEZZAFERRI
Fig. 4.
Lithological
log
and
distribution
of
benthic
foraminifers
in
the
Satskhenisi
section.
N
1
S
1
=
Volhynian;
N
1
S
2
=
Bessarabian.
425
HAUERINIDAE SCHWAGER – TAXONOMIC REVISION OF BESSARABIAN MILIOLIDS (EASTERN PARATETHYS)
Fig. 5. Varidentella reussi costulata. 1a—c – Holotype N, 14/M; 1a—b = side view; 1c = apertural view; drawing after Maisuradze (1971).
2a—c – Paratype N, 14a/M; 2a—b = side view; 2c = apertural view. 3a—c – Specimen N, K/040/127; 3a—b = side view; 3c = apertural view.
T y p e l o c a l i t y : In the gorge of river Ochkhamuri (West-
ern Georgia).
T y p e l e v e l : Bessarabian.
P a r a t y p e s a n d o t h e r m a t e r i a l : Number 14a/M
(Fig. 5.2a—c) is stored at the Georgian National Museum, L.
Davitashvili Institute of Paleobiology (collection of Mai-
suradze), it was found in sediments from the gorge of the riv-
er Ochkhamuri (Western Georgia). The specimen K/040/127
(Fig. 5.3a—c) is stored at the A. Djanelidze Institute of Geol-
ogy (collection of Koiava, Spezzaferri), it was found in
sediments from the gorge of the river Satskhenisi (Eastern
Georgia).
426
MAISURADZE, KOIAVA and SPEZZAFERRI
D e r i v a t i o n o f t h e n a m e : Because of the presence of
striae and costae as ornamentation of the wall texture.
D e s c r i p t i o n : The test is rounded, rarely broadly-oval,
inflated, with a rounded basis and an oblique aperture. The
arrangement of the chambers is crypto-quinqueloculiniform.
The peripheral margin is rounded. Length and width are ap-
proximately equal. The side where 4 to 3 chambers in the
last whorl are observed is always convex. The side with 3
chambers in the last whorl is flat, concave or convex; the su-
tures are depressed and distinct. The aperture is oblique,
elongate and oval, encompassing the whole width of the last
chamber and is bordered by a lip-like structure. The tooth in
the aperture is low and wide, slightly bifid. The wall is thin,
appearing almost translucent and the surface is covered with
dense and fine longitudinal striae.
D i m e n s i o n s : Width – 0.45—0.65 mm; length – 0.32—
0.49 mm; thickness – 0.23—0.35 mm.
V a r i a b i l i t y : The morphological features that character-
ize this subspecies are generally consistent. However, the
size of the tooth, the thickness of the lip-like structure
around the aperture, the rounded to wide oval outline of the
test and the visible number of chambers may vary slightly
from specimen to specimen.
R e m a r k s : The generic attribution of this species to Vari-
dentella is based on the shape of the tooth. It differs from
Varidentella pseudocostata (Venglinski) described by Pope-
scu (1995) in Volhynian sediments from Romania, because
V. reussi costulata possesses a more inflated test, and the
striae ornamenting the test are denser, thinner and less
marked. The specimen described in Cicha et al. (1998) from
Sarmatian sediments from Poland, is more similar to the typ-
ical V. reussi, with an overall smooth wall texture with very
weak and rare striae only along the peripheral margins. This
form is characteristic for silty sediments (Figs. 3 and 4) that
were probably deposited in low energy environments. Hansen
& Dalberg (1979), showed that the presence of pits and costae
in symbiont bearing miliolids such as Amphisorus hemprichii
(Ehrenberg) and Triloculina rupertiana (Brady) indicate that
these features favour the uptake of CO
2
. However, a clear rela-
tionship between ecological conditions and test ornamenta-
tions is difficult to prove for the described species.
D i s t r i b u t i o n : The analysis of our material from several
locations suggests that V. reussi costulata is characteristic
for the Bessarabian and it is present in Georgia, but also in
other Paratethyan regions such as Crimea, Moldova and Bul-
garia (collection Maisuradze).
M a t e r i a l : 20 well preserved specimens.
Suborder: Miliolina Delage & Hérouad 1896
Superfamily: Miliolacea Ehrenberg, 1839
Family: Hauerinidae Schwager, 1876
Subfamily: Hauerininae Schwager, 1876
Genus: Varidentella Łuczkowska, 1972
Varidentella luczkowskae sp. nov.
Fig. 6.1a—c,2a—c,3a—c
1971 Quinqueloculina aff. sartaganica Maisuradze, p. 50, pl. III,
fig. 5a,b,v
M o t i v a t i o n : Karrer (1877) described Quinqueloculina
sarmatica and its variations – var. typica, var. elongata,
var. virgata from Lower Sarmatian deposits of Austria.
Łuczkowska (1972) subsequently attributed Quinqueloculi-
na sarmatica and related forms to the genus Varidentella. In
particular, she also documented Varidentella aff. sartagani-
ca as a species related to V. sartaganica (Krashenninikov).
This latter species is probably the ancestor of many Vol-
hynian and Bessarabian species of Varidentella, e.g. V. reus-
si, V. nanae, V. nanae megrelic (Maisuradze 1971).
Varidentella aff. sartaganica was also described by Mai-
suradze (1971) as possessing a broadly-oval or oval test
(length 1.5 times- or twice its width); a proximal edge, round-
ed and elongated towards the aperture and bordered by a col-
lar-like structure. According to Maisuradze (1971) this species
differs from Varidentella reussi by the more convex proximal
part, the more rounded periphery and the lower neck and
therefore it is now described as Varidentella luczkowskae.
H o l o t y p e : It is presented in Fig. 6.1a—c, and it is stored
at the Georgian National Museum, L. Davitashvili Institute
of Palebiology with the reference number 42/M.
T y p e l o c a l i t y : In the gorge of the river Ochkhamuri
(Western Georgia).
T y p e l e v e l : Bessarabian.
P a r a t y p e s a n d o t h e r m a t e r i a l : Number 42a/M
(Fig. 6.2a—c), is stored at the Georgian National Museum, L.
Davitashvili Institute of Paleobiology (collection of Mai-
suradze) and was found in the gorge of the river Ochkhamuri.
The specimen K/039/127 (Fig. 6.3a—c) is stored at the A.
Djanelidze Institute of Geology (collection of Koiava, Spezza-
ferri), it was found in sediments from the gorge of the river
Satskhenisi (Eastern Georgia).
D e r i v a t i o n o f t h e n a m e : After Eva Łuczkowska be-
cause of her valuable work on miliolids.
D e s c r i p t i o n : The test is oval, slightly convex. The
length is twice the width. Chambers are arranged in quinque-
loculinid-coiling mode. The peripheral margin is rounded.
On one side, 3 to 4 chambers are visible, on the other side,
which is flatter, only 2 to 3 chambers are visible. The cham-
bers are tubular, inflated, arched, larger at the proximal edge
and narrower towards the aperture. The sutures are broad and
distinct. The aperture is semicircular, oblique and elongated
and follows the outline of the chambers. A semicircular, con-
cave and lamellar tooth is present in the aperture. The wall
texture is smooth.
D i m e n s i o n s : Width – 0.37—0.28 mm; length – 0.67—
0.58 mm; thickness – 0.26—0.20 mm.
V a r i a b i l i t y : The characteristic morphology of this spe-
cies is generally consistent, no strong variability has been
observed. However, test outline may be oval to wide oval;
the tooth may display changes in size; visible chambers may
vary in number.
R e m a r k s : This species differs from the related species,
such as Varidentella sartaganica (Krashenninikov) in having
a broader oval test; a more rounded proximal edge at one side,
with tall neck elongated towards the aperture and a large test.
D i s t r i b u t i o n : Bessarabian deposits of Georgia.
M a t e r i a l : Thirty well preserved specimens and twenty
damaged specimens.
427
HAUERINIDAE SCHWAGER – TAXONOMIC REVISION OF BESSARABIAN MILIOLIDS (EASTERN PARATETHYS)
Fig. 6. Varidentella luczkowskae. 1a—c – Holotype N, 42/M; 1a—b = side view; 1c = apertural view; drawing after Maisuradze (1971).
2a—c – Paratype N, 42a/M; 2a—b = side view; 2c = apertural view. 3a—c – Specimen N, K/039/127; 3a—b = side view; 3c = apertural view.
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MAISURADZE, KOIAVA and SPEZZAFERRI
Fig. 7. Affinetrina voloshinovae eldarica. 1a—c – Holotype N, 41/M; 1a—b = side view; 1c = apertural view; drawing after Maisuradze
(1971). 2a—c – Paratype N, 41a/M; 1a—b = side view; 1c = apertural view. 3a—c – Paratype N, K/035/127; 3a—b = side view; 3c = aper-
tural view.
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HAUERINIDAE SCHWAGER – TAXONOMIC REVISION OF BESSARABIAN MILIOLIDS (EASTERN PARATETHYS)
Suborder: Miliolina Delage & Hérouad, 1896
Superfamily: Miliolacea Ehrenberg, 1839
Family: Hauerinidae Schwager, 1876
Subfamily: Hauerininae Schwager, 1876
Genus: Affinetrina Łuczkowska, 1972
Affinetrina voloshinovae eldarica subsp. nov.
Fig. 7.1a—c, 2a—c, 3a—c
1956 Miliolina pseudovoloshinovae Pobedina – Pobedina et al.,
p. 112, pl. XII, fig. 3a,b,v. emended
1971 Quinqueloculina aff. voloshinovae (Bogdanowicz), Mai-
suradze, p. 47, pl. VII, figs. 2a,b,v; 3a,b,v
no 1995 Affinetrina ex group voloshinovae (Bogdanowicz), Popescu,
p. 90, pl. 1, figs. 12—14
M o t i v a t i o n : Affinetrina voloshinovae eldarica, origi-
nally termed Quinqueloculina aff. voloshinovae Bogdanow-
icz was described by Maisuradze (1971) from Bessarabian
deposits of Western Georgia. Similar forms are also present
in coeval deposits of Eastern Georgia. In particular, they oc-
cur in the bore-hole Eldari-1 and in the outcrops Eldari and
Satskhenisi (Koiava 2006). These forms were originally de-
scribed by Pobedina et al. (1956) as Miliolina pseudovoloshi-
novae. However, after a careful revision of the type material
we propose to emend the species Miliolina pseudovoloshino-
vae and include these forms characterized by knob-like orna-
mentation instead of ribs and a chink-like aperture with
elongate tooth in the Q. voloshinovae (Bogdanowicz) group.
H o l o t y p e : It is presented in Fig. 7.1a—c, and it is stored
at the Georgian National Museum. L. Davitashvili Institute
of Paleobiology with the reference number 41/M.
T y p e l o c a l i t y : The gorge of the river Ochkhamuri
(Western Georgia).
T y p e l e v e l : Bessarabian.
P a r a t y p e s : Number 41a/M (Fig. 7.2a—b) is stored at the
Georgian National Museum, L. Davitashvili Institute of Pa-
leobiology (collection of Maisuradze). Number K/035/127
(Fig. 7.3a—c) is stored at the A. Djanelidze Institute of Geo-
logy (collection of Koiava, Spezzaferri). All paratypes were
found in sediments of the gorge of the river Ochkhamuri
(Western Georgia).
D e r i v a t i o n o f t h e n a m e : According to the geograph-
ic location.
D e s c r i p t i o n : The test is irregularly elongated and oval,
more or less convex and angular-rounded at the basis. The
aperture is straight or oblique-truncated. The test is longer
than wide (2 or 2.5 times). The arrangement of chambers is
quinqueloculiniform. The apertural side is rounded to subtri-
angular. The peripheral margin is round. The side displaying
three to four chambers is generally convex. The opposite
side showing two chambers is flat. Chambers are tubular, ob-
lique at the base, especially in the last one. The middle part
of chambers is less wide than at the bottom, and aperture.
The sutures are depressed but not always clearly visible.
The aperture is narrow, straight or oblique, elongated or
elongated-oval, generally wider at one side, with parallel
edges and bordered by a lip. It has a single elongate narrow
and club-shaped tooth. Sometimes the tooth is extended out-
side the aperture.
The wall texture is thick, massive and is covered by nu-
merous irregular longitudinal pustule-like striae, which are
more developed at the bottom of the test where quite often
they are presented by sharp-tipped knobs. Near the end of
the aperture the wall texture becomes smoother.
D i m e n s i o n s : Length – 0.8—0.95 mm; width – 0.36—
0.4 mm; thickness – 0.25—0.35 mm.
V a r i a b i l i t y : The morphological features characterizing
this subspecies are generally consistent. However, the out-
line of the test varies from elongated to slightly oval, the size
of the tooth and the wall ornamentations can also be more or
less pronounced.
R e m a r k s : This subspecies is very similar to Affinetrina
voloshinovae (Bogdanowicz), but it differs in ornamenta-
tion: more pustule-like in A. voloshinovae eldarica and real
striae in A. voloshinovae (Bogdanowicz). Additionally, A.
voloshinovae pecteniformis (Bogdanowicz) differs from this
subspecies in having a hook-like thick low end of the last
chamber, which is sharply projected over the edges of the
test; a wider round base; and a very coarse ornamentation, a
different outline of the aperture and tooth.
D i s t r i b u t i o n : Bessarabian of Georgia and Azerbaijan
(Vangengeim et al. 1989).
M a t e r i a l : Ten well preserved specimens.
Conclusion
Three new Bessarabian miliolid taxa occurring in Georgia
(Eastern Paratethys) are described. They are: Varidentella
luczkowskae, Varidentella reussi (Bogdanowicz) subsp. cos-
tulata, and Affinetrina voloshinovae (Bogdanowicz) subsp.
eldarica. For the first time the classification of Łuczkowska
(1972), based on the morphology of the aperture and the
shape and size of the tooth, is applied to Eastern Paratethyan
miliolids, providing the framework for possible taxonomic
comparison between eastern, central and western taxa.
Acknowledgments: The authors thank the Swiss National
Science Foundation SCOPES Project Ref. IB7320-110693/1
and the European Union INTAS Project No. 1000017-8930
that made the collaboration between Georgia and Switzer-
land possible. Thanks also to all the members of the two
projects for fruitful discussions. The suggestions of the re-
viewers (S. Filipescu and an anonymous) greatly improved
the quality of this article. The photos of the foraminifera
were obtained at the Department of Geosciences, University
of Fribourg. Thanks to C. Neururer (Fribourg) for his help
operating the SEM. A warm thank to R. Waite (Fribourg) for
correcting the English form.
References
Bogdanowicz A.K. 1947: The results of study of Miocene foramin-
ifera of Crimea-Caucasus region. In: Microfauna of oil loca-
tions of Caucasus, Emby and Middle Asia. Trans. VNIGRI
5—34 (in Russian).
Bogdanowicz A.K. 1952: Miliolidae and Peneroplidae the fossil
430
MAISURADZE, KOIAVA and SPEZZAFERRI
foraminifera of USSR. Trans. VNIGRI 64, 1—338 (in Russian).
Bogdanowicz A.K. 1959: The basis of paleontology. The Protero-
zoa. Acad. Sci. USSR, Moscow, 1—36 (in Russian).
Bogdanowicz A.K. 1960: About new and little known foraminifers
from the Miocene of Western Pre-Caucasus. Trans. VNIGRI 3,
241—263 (in Russian).
Bogdanowicz A.K. 1981: The classification of small foraminifera
of Mezo-Cenozoic. Trans. VNIGRI No. 3, 3—127 (in Russian).
Brady H.B. 1884: Report on the Foraminifera collected by H.M.S.
Challenger during the years 1873—1876. In: Thomson C.W. &
Murray J. (Eds.): Report of the scientific results of the voyage
of H.M.S. Challenger during the years 1873—76 under the com-
mand of Captain George S. Nares and Captain Frank Tourle
Thomson. London, Edinburgh, Dublin, IX, 1—814.
Cicha I., Rögl F., Rupp C. & Čtyroká J. 1998: Oligocene—Miocene
foraminifera of the Central Paratethys. Abh. Senckenberg.
Naturforsch. Gesell. 549, 1—325.
Delage Y. & Hérouad E. 1896: Traité de Zoologie Concr
e
te. Le
Cellule et les Protozoaires. Schleichter Fr
e
res, Paris, 1, 1—589.
d’Orbigny A. 1826: Tableau méthodique de la classe des Céphalop-
odes. Ann. Sci. Natur. 7, 245—314.
Ehrenberg G.C. 1839: Über die Bildung der Kreidefelsen und des
Kreidemergels durch unsichtbare Organismen. Phys. Abh.
Kön. Akad. Wiss. Berlin 1838 (1840: separate 1839), 59—147.
Finlay J.A. 1947: New Zealand Foraminifera: key species in stratig-
raphy. New Zealand J. Sci. Tech. 28, 5 (sec. B), 272—275.
Hansen J.H. & Dalberg P. 1975: Symbiotic algae in milioline fora-
minifera: CO
2
uptake and shell adaptation. Bull. Geol. Soc.
Denmark 28, 47—55.
Harzhauser M. & Piller W.E. 2007: Benchmark data of a changing
sea – palaeogeography, palaeobiogeography and events in the
Central Paratethys during the Miocene. Palaeogeogr. Palaeo-
climatol. Palaeoecol. 253, 8—31.
Iaccarino S. 1985: Mediterranean Miocene and Pliocene planktic
foraminifera. In: Bolli H.M., Saunders J.B. & Perch-Nielsen
K. (Eds.): Plankton stratigraphy. Cambridge Earth Sci. Ser.,
Cambridge Univ. Press, 283—314.
Kamladze G. & Kachkachishvili M. 1976: Summarizing of geolog-
ic-geophysical materials on Abkhazia and Megrelia. Rep. Abk-
hazian-Megrelian Thematic Geol. Party, Tbilisi, 1—216 (in
Russian).
Karrer F. 1877: Geologie der Kaiser Franz-Josephs Hochquellen-
Wasserleitung. Eine Studie in den Terliar-Bildungen am We-
strande des alpinen Tiles der Niederung Wien. Abh. K.K. Geol.
Reichsanst. Wien 9, 1—420.
Koiava K. 2006: The biostratigraphy of Sarmatian deposits of East-
ern Georgia on the base of Foraminifera. PhD Thesis, Alexan-
dre Djanelidze Inst. Geol., Tbilisi, 1—163 (in Georgian).
Kolesnikov V. 1935: Sarmatian Mollusks. Paleontology USSR, Mos-
cow, 1—507 (in Russian).
Lamarck J.B. 1804: Suite des mémoires sur les fossiles desenvirons
de Paris. Ann. Mus. Nat. Hist. Natur. 5, 349—357.
Loeblich A.R. Jr. & Tappan H. 1964: Sarcodina chiefly “The camoe-
bians” and Foraminifera. In: Moore R.C. (Ed.): Treatise of in-
vertebrate paleontology. C. Protista, New York, 2, 1, 1—510.
Loeblich A.R. Jr. & Tappan H. 1988: Foraminiferal genera and their
classification. Van Nostrand Reinhold, New York, 1—2, 1—970.
Łuczkowska E. 1972: Miliolidae (Foraminiferida) from Miocene of
Poland//part I. Revision of the classification. Acta Palaeont.
Pol. XVII, 3, 342—377.
Łuczkowska E. 1974: Miliolidae (Foraminiferida) from Miocene of
Poland//part II. Biostratigraphy, palaeoecology and systemat-
ics. Acta Palaeont. Pol. XIX, 1, 1—176.
Maisuradze L.S. 1971: The Foraminifera of Western Georgia Sar-
matian. Tbilisi, 1—120 (in Russian).
Papava D. 1974: Geological structure and oil-and-gas bearing of the
middle flow of the river Mtkvari. Report of Adjara-Trialety
Thematic Geologic Party, Tbilisi, 1—254 (in Russian).
Piller W.E. & Harzhauser M. 2005: The myth of the brackish Sar-
matian Sea. Terra Nova 17, 450—455.
Pobedina V.M., Voroshilova A.T., Rybina O.I. & Kuznetsova T.V.
1956: The reference book on microfauna of middle and upper
Miocene deposits of Azerbaijan. Baku, 3—179.
Popescu G. 1995: Contribution to the knowledge of the Sarmatian
Foraminifera of Romania. Roman. J. Paleontology 76, 85—98.
Popov S.V., Akhmetev M.A., Zaporozhets N.I., Voronina A.A. & Stol-
yarov A.S. 1993: Evolution of Eastern Paratethys in the Late
Eocene—Early Miocene. Stratigr. Geol. Correlation 1, 6, 572—600.
Rögl F. 1985: Late Oligocene and Miocene planktic foraminifera of
the Central Paratethys. In: Bolli H.M., Saunders J.B. & Perch-
Nielsen K. (Eds.): Plankton stratigraphy. Cambridge Earth Sci.
Ser., Cambridge Univ. Press, 315—328.
Rögl F. 1999: Mediterranean and Paratethys. Facts and hypothesis
of an Oligocene to Miocene paleogeography (short overview).
Geol. Carpathica 50, 339—349.
Rögl F. 2001: Mid-Miocene circum-Mediterranean paleogeography.
Berichte Inst. Geol. Paläont., K.-F.-Univ. Graz 4, 49—59.
Schlumberger C. 1887: Note sur le genre Planispirina. Bull. Soc.
Zool. France 12, 105—118.
Schlumberger C. 1893: Monographie des Miliolidées du Golfe de
Marseille. Mém. Soc. Zool. France 6,199—224.
Schwager C. 1876: Saggio di una classificazione dei foraminiferi
avuto riguardo alle loro famiglie naturali. Boll. Comitato Geol.
Ital. 8, 18—153.
Spezzaferri S. & Ćorić S. 2001: Ecology of Karpatian (Early Mi-
ocene) foraminifera and calcareous nannoplankton from Laa an
der Thaya, Lower Austria: a statistical approach. Geol. Car-
pathica 200, 361—374.
Steininger F.F. & Wessely G. 2000: From the Tethyan Ocean to the
Paratethys Sea: Oligocene to Neogene stratigraphy, paleogeog-
raphy and paleobiogeography of the circum-Mediterranean re-
gion and the Oligocene to Neogene Basin evolution in Austria;
Aspects of Geology in Austria. Mitt. Österr. Geol. Gesell. 92,
95—116.
Vangengeym E.A., Gabuniya L.K., Pevzner M.A. & Tsiskarishvili
G.V. 1989: Stratigraphical position of Transcaucasus Hippari-
on fauna occurrences in light of magnetic stratigraphy data.
Trans. Acad. Sci. USSR, Ser. Geol. 8, 70—78 (in Russian).
Vella P. 1957: Foraminifera from Cook Strait. In: Studies in New
Zealand Foraminifera, Part 1. Bull. Paleont. New Zealand
Geol. Surv. 28, 1—41.
Wiesner H. 1931: Die Foraminiferen der deutschen. Südpolar –
Expedition 1901—1903. Drygalski’s Deutsche Südpolar Exp.,
Zool., Berlin-Leipzig 20, 53—165.
Williamson W.C. 1858: On the recent Foraminifera of Great Brit-
ain. Roy. Soc. Publ., London, 1—107.
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