GEOLOGICA CARPATHICA
, APRIL 2017, 68, 2, 109 –118
doi: 10.1515/geoca-2017-0009
www.geologicacarpathica.com
Systematic updates of the agglutinated foraminiferal genus
Colominella Popescu, 1998: insights from sectioned specimens
NICOLETTA MANCIN
1,
and MICHAEL A. KAMINSKI
2
1
Dipartimento di Scienze della Terra e dell’Ambiente, Università degli Studi di Pavia, via Ferrata 1, 27100 Pavia, Italy;
nicoletta.mancin@unipv.it
2
Geosciences Department, King Fahd University of Petroleum & Minerals, PO Box 701 KFUPM, Dhahran, 31261, Saudi Arabia;
kaminski@kfupm.edu.sa
(Manuscript received July 7, 2016; accepted in revised form November 30, 2016)
Abstract: The occurrence of agglutinated foraminiferal specimens belonging to the Badenian (middle Miocene) genus
Colominella Popescu, 1998 was recently documented for the first time in a lower Pliocene succession of the western
Mediterranean area. Direct comparisons with topotype specimens of Colominella paalzowi (Cushman 1936), sampled in
the Badenian type section of Lăpugiu de Sus (Transylvania), show that the Pliocene individuals from the western
Mediterranean morphologically resemble the type species C. paalzowi, but they also differ in possessing a longer biserial
chamber arrangement with a higher number of internal chamber partitions, in lacking a clear early triserial stage and
in having a more complex microstructure of the agglutinated wall, thereby supporting the idea that the Pliocene
Mediterranean specimens represent a new, more highly evolved species. The fact that the Pliocene individuals from the
Mediterranean appear to be more evolved with respect to the Badenian specimens from Paratethys represents an interesting
evolutionary development of the genus Colominella that also permits the known stratigraphical and geographical range
of the genus, previously limited to the middle Miocene (Badenian) of the Paratethys, to be extended.
Keywords: Mediterranean Pliocene, Badenian Paratethys, agglutinated foraminifera, taxonomy, wall microstructure,
SEMEDS analysis.
Introduction
The genus Colominella Popescu, 1998 has been formally
described from the Paratethyan Badenian (middle Miocene)
Kostej succession, cropping out in Transylvania (Popescu et
al. 1998; Kaminski 2004) and is based on a species (type spe
cies Textulariella paalzowi) first studied by Cushman (1936)
from the same locality. The type species was subsequently
recorded in the Badenian of the Rauchstallbrunngraben in the
Vienna Basin (Popescu et al. 1998), in the lower Badenian
Wagna succession (Styrian Basin) in the western portion of the
Pannonian basin system within the AlpineCarpathian arch
(Spezzaferri et al. 2004), and more recently, in the lower Plio
cene record of the Albenga Basin in the western Mediterra
nean region (Mancin et al. 2012).
According to Cushman’s original description, Colominella
(ex Textulariella) paalzowi is characterized by a very large
(4 mm long and 1 mm in diameter), conical, mostly biserial
test, triserial in the early stage, with typical internal vertical
partitions, comparatively few in number, and a canaliculate
test wall with a coarsely agglutinated surface.
Colominella likely evolved from the genus Matanzia (which
is also canaliculate) during the Oligocene to middle Miocene
(Kaminski & Cetean 2011). Colominella and Matanzia have
been recently placed in the subfamily Colominellinae Popescu,
1998 together with two additional closelyrelated genera:
Colomita GonzalesDonoso, 1968 and Cubanina Palmer,
1936 (Kaminski & Cetean 2011 and references therein). In
spite of this recent systematic review of the genus, not much is
known about the test wall microstructure of Colominella nor
about its stratigraphical and geographical range, which is cur
rently reported as limited to the Badenian of Paratethyan suc
cessions (Kaminski & Cetean 2011 and references therein).
In this work, we used high definition SEMEDS images of
entire and sectioned specimens, to investigate both the mor
phology and the test wall microstructure of some agglutinated
foraminifera belonging to the genus Colominella coming from
two sections exposed in the western Mediterranean area (Rio
Torsero section) and in the Paratethyan Transylvanian Basin
(Lăpugiu de Sus section) (Fig. 1A’). The main purpose of this
study is to compare the Pliocene Mediterranean individuals
with the Badenian topotypes in order to verify whether or not
the Pliocene Mediterranean specimens described by Mancin et
al. (2012) were correctly assigned to the species Colominella
paalzowi, thereby extending the known stratigraphical and
geographical range of the genus.
Materials and methods
The Lăpugiu de Sus section crops out in one of the most
famous fossiliferous sites of the Transylvanian Basin (Hune
doara county, Rumania) along the Mureş River (Fig. 1A’). The
very rich mollusc fauna, in particular bivalvia and gastropods,
was mentioned in 1863 by Stur and then was exhaustively
studied, as testified by the numerous palaeontological and
stratigraphical data subsequently published (e.g., Petrescu et
al. 1990; Chira 1994; Chira et al. 2000; Chira & Voia 2001;
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Harzhauser et al. 2003; Tămaş et al. 2013). The Badenian fora
miniferal assemblages from the same locality were the subject
of numerous studies since the pioneering work of Cushman
(1936) (e.g., Nicorici et al. 1994; Popescu & Crihan 2005;
Krézsek & Filipescu 2005; Báldi et al. 2002; Báldi 2006;
Filipescu & Silye 2008; Boga 2013; de Leeuw et al. 2013).
These studies provided a complete biostratigraphical and envi
ronmental framework for the central Paratethyan area.
In the western Mediterranean region, the Rio Torsero section
crops out along the homonymous rivulet in the easternmost
portion of the Albenga basin (western Liguria; Italy) (Fig. 1A’).
This Pliocene outcrop has been well known since the 20
th
cen
tury thanks to its rich mollusc faunas (mainly gastropods) as
documented by several studies (e.g., Bernasconi & Robba 1984,
1994; Solsona 1999; Solsona & Martinell 1999; Andri et al.
2005; Harzhauser & Kronenberg 2008). Published micro
palaeontological data (Violanti 1987; Mancin et al. 2012)
pointed out the depositional environment of the Rio Torsero
section, which probably recorded the sedimentation in the
circalittoral zone, close to the shelf edge.
For the present study, we examined, under a Scanning Elec
tron Microscope (SEM) equipped with an Xray Energy
Dispersive Spectrometer (EDS), four topotype specimens of
Colominella paalzowi, that were isolated and picked from
samples collected from the Badenian Lăpugiu de Sus type sec
tion (Fig. 1A’). In order to directly compare the Paratethyan
Fig. 1. A–A’: Simplified geographical map of the Mediterranean area and Eastern Europe with the location of the studied sections (1 — Lăpugiu
de Sus section in Romania; 2 — Rio Torsero section in Italy). B–C: Schematic palaeogeographical maps of the Africa–Eurasia collision zone
reconstructed for the Early Pliocene (partly modified and redrawn after Popov et al. 2006) and the Middle Miocene (partly modified and
redrawn after Rögl 1999). Note that during the Badenian (Langhian) the Central Paratethys was connected with the Mediterranean Sea favouring
the great similarity of their marine palaeontological record (de Leeuw et al. 2013). Since the late Badenian (early Serravallian) the connection
failed causing the progressive faunal differentiation in the two basins (de Leeuw et al. 2013).
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Fig. 2. Morphological variability of some Colominella specimens from the Pliocene Rio Torsero (images 1–8) and the Badenian Lăpugiu
de Sus (images 9–11) sections. SEM images are in secondary electrons (SE).
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Fig. 3. Images at the stereomicroscope of a topotype specimen of
Colominella paalzowi from the Badenian Lăpugiu de Sus section
showing the typical conical shape (1a), a clearly evident early triserial
stage without internal partitions (arrow in 1b) and a rounded profile in
apertural view (1c).
and Mediterranean specimens of Colominella, we also ana
lysed five new specimens of Colominella isolated and picked
from the same samples previously studied by Mancin et al.
(2012) collected from the lower Pliocene Rio Torsero section
(Fig. 1A’). Moreover, to better compare new results with data
reported in the previous paper of Mancin and coauthors, all
the collected specimens were prepared and analysed following
the same methodology described in Mancin et al. (2012) and
recently standardized by Mancin et al. (2014). The studied
specimens are stored in the micropalaeontological collection
of the Department of Earth and Environment Sciences of the
University of Pavia (Italy).
Results
Morphology of the Colominella test
The Pliocene Mediterranean specimens have a large test
(from 2.5 to 3 mm long and 1.5 to 2 mm wide) with an elongate
morphology typically with a welldeveloped biserial stage, but
lacking a clear early triserial stage (Fig. 2, images 1–5).
The internal radial partitions forming alcoves appear quite soon
during the test growth, sharply increasing in number passing
from the earlier to the later chambers (Fig. 2, images 6–8).
The Badenian topotypes are smaller (1 to 1.5 mm long and
0.5 to 1 mm wide), with a characteristic conical and more fusi
form shape due to the chambers that sharply increase in size
during ontogeny (Fig. 2, images 9–11; Fig. 3) and the occur
rence of a welldeveloped early triserial stage (Fig. 3, image
1b). Chamber partitions appear later in the ontogenetic growth
as they are missing from the earlier triserially arranged cham
bers (Fig. 3, image 1b arrow). The number of vertical parti
tions forming alcoves is also different in the Pliocene and
Badenian individuals: alcoves are more numerous in Pliocene
specimens (7 to 8 in the last chambers; Fig. 2, image 8) while
they are less numerous (from 4 to 5) in the Badenian individuals
(Fig. 2, image 9; see also Spezzaferri et al. 2004, Pl.6, fig. 9).
The Pliocene Mediterranean specimens, moreover, have
an external, homogeneous smooth surface; a sort of external
pavement that covers the open entrances of the canaliculi
crossing the test wall. In some portions of the test, this pave
ment is missing because it was partly abraded by erosion
(Fig. 2, images 1–4). Conversely, in the Badenian specimens
the test surface is more coarsely agglutinated and made of
heterogeneous grains consisting also of the remains of small
calcareous foraminifera (Fig. 2, images 10 –11a).
Microstructure of the agglutinated wall
In sectioned specimens, the microstructure of the aggluti
nated wall appears quite different in the Pliocene individuals
with respect to the Badenian specimens. In the former, the test
wall is thicker (over 100 mm) and abundantly crossed by cana
liculi (Fig. 4, images 1a–c, 2a–b, 3a–b); they are straight and
radial in the inner portion of the wall but become branching in
the outer part (Fig. 4, images 1b, 1c, 2b). Moreover, canaliculi
are present mostly in the upper portion of alcoves towards the
external margin and are missing internally and on the secon
dary septa (Fig. 4, image 1d). The agglutinated granules are
selected with respect to their size and disposition within the
test wall: the largest grains are positioned toward the outside
margin, while the smallest ones are internally arranged, simu
lating a sort of layered microstructure of the agglutinated wall
(Fig. 4, images 1a, 1d, 2b, 3a–b). Where the agglutinated
wall is crossed by canaliculi, the coarsegrained particles are
embedded in an aggregate of very small detrital granules, 3 to
5 mm in size (Fig. 4, image 1b white arrow), probably to favour
the development of the canaliculi themselves. The grain selec
tivity persists unchanged throughout the whole test (Fig. 4,
images 1, 2, 3).
In the Badenian specimens of C. paalzowi the test wall is
less thick (ca. 90 mm or less) and abundantly crossed by cana
liculi that, as in the Pliocene individuals, are present mostly in
the upper portion of alcoves towards the external margin and
are missing on the secondary septa (Fig. 5, images 1a, 2a, 3b).
However, the agglutinated granules are not always selected in
size and disposition (Fig. 5, images 1a–b, 3a–b) resulting in
a more disorganized wall microstructure without a distinctly
layered grain arrangement.
Fig. 4. Sectioned specimens of Colominella from the Pliocene Rio Torsero section. SEM images are in backscattered electrons (BSE).
1: Specimen (the same of Fig. 2, image 1) longitudinally sectioned to show the internal partition of the chambers and the microstructure of the
agglutinated wall; 1a — detail of the agglutinated wall crossed by canaliculi,1b — enlargement of “a” showing how the coarsegrained
particles are embedded by numerous small detrital granules (white arrow), 1c–d — microstructural details of both the test wall and a septum
showing the grains which are selected in terms of size and disposition, forming a layered microstructure (arrow). Note that only the test wall is
crossed by canaliculi that are missing on the septa. 2: Specimen (the same of Fig. 2, image 3) longitudinally sectioned; 2 a–b — details of the
agglutinated wall. 3: Specimen (the same of Fig. 2, image 5) longitudinally sectioned; 3 a–b — details of the agglutinated wall. Note that in all
the sectioned specimens the microstructural characteristics of the agglutinated wall persist unchanged during the test growth.
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Chemical-mineralogical composition of the agglutinated
grains
Even from a compositional point of view the Pliocene and
Badenian specimens are different. The former ones are charac
terized by agglutinated grains that are compositionally more
homogeneous and mostly made of dolomite, quartz, with
a minor amount of calcite and Kfeldspar (Fig. 6, image 1a).
Quartz grains mostly form the external pavement that covers
the open entrance of canaliculi (Fig. 6, image 1d Simap),
on the other hand, the internal grains are mostly of dolomite.
All the grains are cemented by calcite cement that appears as
cloudy light grey areas among the detrital grains (Fig. 4,
images 1c–d, 2a–b, 3a–b; Fig. 6, image 1c Camap).
Conversely, in the Badenian specimens, the agglutinated
grains are compositionally more heterogeneous and formed by
calcite (Fig. 7, images 1c, 2c Camaps), quartz (Fig. 7, images
1a, 2a Simaps), with a minor amount of dolomite and albite
(Fig. 7, images 1b, 2b Mgmaps and 2d Namap). It is note
worthy that the quartz grains are heterogeneously distributed
within the test wall even within the internal septa (Fig. 7,
images 1a and 2a Simaps) and they do not concentrate
towards the external margin of the test wall to form a pave
ment covering the canaliculi. Moreover, some agglutinated
grains are remains of planktonic foraminifera that, on the con
trary, are never observed in the Pliocene specimens (Fig. 2
image 11a; Fig. 5, image 1a).
Do the Pliocene Mediterranean specimens belong
to the type species C. paalzowi?
The direct comparison between the specimens of Colomi-
nella from the Pliocene Mediterranean record and the topo
type individuals of C. paalzowi from the lower Badenian
Paratethyan Lăpugiu de Sus section shows that the Pliocene
Mediterranean specimens have a different, more complex
test. The complexity of the agglutinated test, in terms of
a major number of internal partitions, a test wall micro
structure with a layered structure and a strong selection of
the agglutinated grains, represents a higher grade of evolution
probably driven by the adaptive response to different ecolo
gical conditions that persisted for a long time, most likely
since the early Serra vallian (ca. 13.5 Ma) when the marine
connection between the Mediterranean and Paratethys failed
(Rögl 1998, 1999) (Fig. 1 A–C), promoting the progressive
and inevitable faunal differentiation in the two basins
(de Leeuw et al. 2013). Mancin et al. (2012) interpreted the
increased complexity of the Colominella test as an evolutio
nary adaptation to perform kleptoplastidy and/or to house
photosymbionts probably at shallow bathymetries in warm
water environments.
The collected results support the idea that the Pliocene
Colominella specimens from the Mediterranean Rio Torsero
section are different from a taxonomic point of view, and
therefore they cannot be assigned to the topotype species
C. paalzowi (Cushman). The Pliocene Mediterranean speci
mens more probably represent a new, younger, more highly
evolved Colominella species.
Our understanding of the occurrence of a new species of
Colominella in the Mediterranean Pliocene record awaits the
formal description of the type species and a formal update
of the stratigraphical and geographical range of the genus
Colominella.
Final remarks
This study reports, through high definition SEMEDS
images of entire and sectioned agglutinated specimens, the
direct comparison between specimens of Colominella sam
pled in a Pliocene succession from the western Mediterranean
previously studied by Mancin et al. (2012) and topotypes of
the species Colominella paalzowi recently sampled in the
Badenian Lăpugiu de Sus type section.
The Pliocene Mediterranean specimens cannot be taxo
nomically attributed to the type species C. paalzowii, but
more probably they represent a new, more highly evolved
species. Further investigations on other middle Miocene–
Pliocene shallow water records from the Mediterranean Sea
are needed to provide a new understanding of how the genus
Colominella had evolved in the Mediterranean Sea since the
early Serravallian, and to formally describe the new Pliocene
species.
Acknowledgements: The authors kindly thank Arvedi’s
Laboratory (CISRiC, University of Pavia) and particularly
E. Basso for SEMEDS analyses. We thank Christian Boga
(formerly from Bucharest University) for providing speci
mens of Colominella paalzowi from Romania. This work was
financially supported by FAR and FRG funds (University of
Pavia) and by the Deanship of Scientific Research, King Fahd
University of Petroleum & Minerals, through grants IN121028
and RG1401.
Fig. 5. Sectioned topotype specimens of C. paalzowi from the Badenian Lăpugiu de Sus type section. SEM images are in backscattered
electrons (BSE). 1: Specimen (the same of Fig. 2, image 9) longitudinally sectioned to show the internal features of the test and
the micro structure of the agglutinated wall. 1 a–b — Details of the agglutinated wall crossed by canaliculi. Note that granules are quite
chaotically distri buted and poorly selected in terms of size and disposition within the wall; note also that some of them are formed by
fossil remains of planktonic foraminifera (arrow). 2: Specimen (the same of Fig. 2, image 10) longitudinally sectioned; 2 a–b — details
of the agglutinated wall. 3: Specimen (the same of Fig. 2, image 11) longitudinally sectioned; 3 a–b — details of the agglutinated wall.
The microstructural characte ristics of the agglutinated wall appear to persist unchanged during test growth in all the sectioned Pliocene
specimens.
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Fig. 7. Elemental characterization of the agglutinated grains in two specimens of C. paalzowi from the Badenian Lăpugiu de Sus type section.
Images are in backscattered electrons (BSE). 1–2: Longitudinally sectioned specimens of C. paalzowi showing the test wall microstructure
with reported the spots for standardless microanalyses (coloured crosses) together with the corresponding EDX spectrum used to composi
tionally characterize the agglutinated grains (in this case a grain of albite). a – d — Elemental maps showing the distribution of Silicon (a),
Magnesium (b), Calcium (c) and Sodium (d) in the whole sectioned tests recorded in images 1 and 2. Mg and Si are here considered as discri
minated elements of dolomite and quartz, respectively, whereas Na is distinctive of albite. Note that most of the agglutinated grains are calcite
while Si is quite rare and does not concentrate only in the external portion of the test but is also internally distributed sometimes in the upper
margin of the septa (arrows). Mg and Na are quite rare and without any particular distribution within the test wall.
Fig. 6. Elemental characterization of the agglutinated grains in one specimen of Colominella from the Pliocene Rio Torsero section. Images are
in backscattered electrons (BSE). 1a — Longitudinally sectioned specimen of Colominella showing a detail of the test wall microstructure (a)
with indication of the spots for standardless microanalyses (coloured crosses) together with the corresponding EDX spectra used to composi
tionally characterize the agglutinated grains. 1b – d — Elemental maps showing the distribution of Magnesium (b), Calcium (c) and Silicon (d)
in the portion of test of Colominella recorded in image 1a. Mg and Si are here considered as discriminated elements of dolomite and quartz,
respectively. Note that Si mostly concentrates in the external portion of the test forming the pavement that covers the canaliculi; on the other
hand, Mg and Ca occur abundantly in the internal portion of the test wall.
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References
Andri E., Tagliamacco A., Testa M. & Marchini A. 2005: Fossil mol
lusk faunas from the Rio Torsero. Nuova Editrice Genovese,
Genova, 1–286 (in Italian).
Báldi K. 2006: Paleoceanography and climate of the Badenian
(middle Miocene, 16.4–13.0 Ma) in the Central Paratethys based
on foraminifera and stable isotope (d
18
O and d
13
C) evidence. Int.
J. Earth Sci. (Geol. Rundsch) 95, 119–142.
Báldi K., Benkovics L. & Sztanó O. 2002: Badenian (Middle
Miocene) basin development in SW Hungary: geohistory based
on quantitative paleobathymetry of foraminifera. Int. J. Earth
Sci. (Geol. Rundsch) 91, 490–504.
Bernasconi M.P. and Robba E. 1984: The Pliocene Turridae from
western Liguria. I Clavinae, Turrinae, Turriculinae, Crassispi
rinae, Borsoniinae, Clathurellinae. Boll. Mus. Reg. Sci. Nat.
Torino 2, 257–358.
Bernasconi M.P. & Robba E. 1994: Notes on some Pliocene Gastro
pods from Rio Torsero, western Liguria, Italy. Riv. It. Paleont.
Stratig. 100, 1, 71–102.
Boga C.R. 2013: The study of the foraminifera and ostracoda micro
fuana of the Badenian deposits from Coşului Valley, Lăpugiu de
Sus, Hunedoara county. In: The 9
th
Romanian Symposium on
Paleontology, Iasi, October 25–26, Abstract Book. Iasi, 7.
Chira C. 1994: Catalogue of the bivalvia collection of Lăpugiu de Sus
section preserved at the PaleontologyStratigraphy Museum in
the University of Cluj. In: The Miocene from the Transilvanian
Basin, Romania. ClujNapoca, 75–80.
Chira C. & Voia I. 2001: Middle Miocene (Badenian) Conidae from
Lăpugiu de Sus, Romania: systematical and paleontological
data. Studia Univ. Babeş-Bolyai, Geol. XLVI, 2, 151–160.
Chira C., Filipescu S. & Codrea V. 2000: Palaeoclimatic evolution in
the Miocene from Transylvanian depression reflected in the
fossil record. Geol. Soc. London, Spec. Publ. 1, 181, 55–64.
Cushman J.A. 1936: New genera and species of the families
Verneuilinidae and Valvulinidae and of the subfamily Virguli
nidae. Cushman Lab. Foram. Res., Spec. Publ. 6, 44.
de Leeuw A., Filipescu S., Maţenco L., Krijgsman W., Kuiper K. &
Stoica M. 2013: Paleomagnetic and chronostratigraphic con
straints on the Middle to Late Miocene evolution of the Transyl
vanian Basin (Romania): implications for Central Paratethys
stratigraphy and emplacement of the TiszaDacia plate. Global
Planet. Change 103, 82–98.
Filipescu S. & Silye L. 2008: New Paratethyan biozones of planktonic
foraminifera described from the Middle Miocene of the Tran
sylvanian Basin (Romania). Geol. Carpath. 59, 6, 537–544.
Harzhauser M., Mandic O. & Zuschin M. 2003: Changes in Para
tethyan marine molluscs at the early/middle Miocene transition:
diversity, palaeogeography and palaeoclimate. Acta Geol.
Polonica 53, 4, 323–339.
Harzhauser M. & Kronenberg G.C. 2008: A note on Strombus
coronatus Defrance, 1827 and Strombus coronatus Röding, 1798
(Mollusca: Gastropoda). The Veliger 50, 2, 1–9.
Kaminski M.A. 2004: The new and reinstated genera of Agglutinated
Foraminifera published between 1996 and 2000. In: Bubík M. &
Kaminski M.A., (Eds): Proceedings of the Sixth International
Workshop on Agglutinated Foraminifera. Grzybowski Found.
Spec. Publ. 8, 257–271.
Kaminski M.A. & Cetean C.G. 2011: The systematic position of the
foraminiferal genus Cubanina Palmer, 1936 and its relationship to
Colominella Popescu, 1998. Acta Paleont. Romania 7, 231–234.
Krézsek C. & Filipescu S. 2005: Middle to Late Miocene sequence
stratigraphy of the Transylvanian Basin (Romania). Tectono-
physiscs 410, 437–463.
Mancin N., Basso E., Pirini C. & Kaminski M.A. 2012: Selective
mineral composition, functional test morphology and paleoeco
logy of the agglutinated foraminiferal genus Colominella
Popescu, 1998 in the Mediterranean Pliocene (Liguria, Italy).
Geol. Carpath. 63, 6, 491–502.
Mancin N., Basso E., Kaminski M.A. & Dogan A.U. 2014: A stan
dard SEMEDS methodology to determine the test microstruc
ture of fossil agglutinated foraminifera. Micropaleontol. 61,
13–26.
Nicorigi E., Bedelean I., Mészáros N. & Petrescu I. 1994: The Mio
cene from the Transylvanian Basin, Romania. Carpathica,
ClujNapoca, 1–224 .
Petrescu E., Meszaros N., Chira C. & Filipescu S. 1990: Lower
Badenian paleoclimate at Lăpugiu de Sus (Hunedoara Country),
on account of paleontological investigation. Studia Univ.Babeş-
Bolyai, Geol. XXXV, 2, 13–22.
Popescu G. & Crihan I.M. 2005: Middle Miocene Foraminifera from
Romania: Order buliminida, part I. Acta Palaeont. Romanie 5,
379–396.
Popescu G., Cicha I. & Rögl F. 1998: Systematic Notes. In: Cicha I.,
Rögl F., Rupp C. & Cytroka J. (Eds): Oligocene–Miocene
Foraminifera of the Central Paratethys. Abhandlungen der
senckenbergischen naturforschenden Gesellschaft 549, 69–325.
Popov S.V., Shcherba I.G., Ilyina LB., Nevesskaya L.A., Paramonova
N.P. Khondkarian S.O. & Magyar I. 2006: Late Miocene to
Pliocene palaeogeography oft he Paratethys and its relation to
Mediterranean. Palaeogeog. Palaeoclim. Palaeoecol. 238,
91–106.
Rögl F. 1998: Palaeogeographic considerations from Mediterranean
and Paratethys seaways. Ann. Natuhist. Mus. Wien. 99A,
279–310.
Rögl F. 1999: Mediterranean and Paratethys. Facts and hypothesis of
an Oligocene to Miocene paleogeography (short overview).
Geol. Carpath. 50, 4, 339–349.
Solsona M. 1999: Systematics and description of the families Tonni
dae, Ficidae and Cassidae (Tonnoidea, Gastropoda) from the
Pliocene of the northwestern Mediterranean. Bull. Inst. Cat.
Hist. Nat. 67, 69–90 (in Italian with English abstract).
Solsona M. & Martinell J. 1999: Protoconch as a taxonomic tool in
Gastropoda systematics. Application in the Pliocene Mediterra
nean Naticidae. Geobios 32, 3, 409–419.
Spezzaferri S., Rögl F., Ćorić S. & Hohenegger J. 2004: Paleoenvi
ronmental changes and agglutinated foraminifera across the
Karpatian/Badenian (early/middle Miocene) boundary in the
Styrian basin (Austria, Central Paratethys). In: Bubík M. &
Kaminski M.A., (Eds): Proceedings of the Sixth International
Workshop on Agglutinated Foraminifera. Grzybowski Found.
Spec. Publ. 8, 423–459.
Stur D. 1863: Bericht uber die geologische Ubersichtaufnahme des
sudvestichen Sieberburgens im Sommer 1860. Jahrb. D. k. k.
Geol. R.A., Wien XII, 33–120.
Tămaş A., Tămaş D.M. & Popa M.V. 2013: Badenian small gastro
pods from Lăpugiu de Sus (Făget Basin, Romania). Roissidae
Family. Acta Palaeont. Romaniae 9, 1, 57–66.
Violanti D. 1987: Taxonomical and paleoenvironmental analyses of
foraminiferal assemblages from the Ligurian Pliocene (Rio
Torsero). Boll. Mus. Reg. Sci. Nat. Torino, 5, 1, 239–293
(in Italian with English abstract).