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Upper Barremian representatives of  Dzirulina Nutsubidze,

1945 (Terebratellidina, Brachiopoda) from eastern Serbia








Institute of Regional Geology and Paleontology, Faculty of Mining and Geology, University of Belgrade, Kamenička 6, P.B. 227,

11 000 Belgrade, Republic of Serbia;  vrad@eunet.yu


Natural History Museum, Njegoševa 51, 11 000 Belgrade, Republic of Serbia;  brad@net.yu

(Manuscript received February 21, 2005; accepted in revised form March 16, 2006)

Abstract: A new Late Barremian terebratellid species of Dzirulina Nutsubidze, 1945 of Bežište, Suva Planina Mountain,
eastern Serbia, which was earlier described by Petković (1930) was found at the same locality as Waldheimia (Zeilleria)
zujovici Antula, 1903. The new species is compared with Dzirulina pseudojurensis (Leymerie), and the variability in the
external features of this widespread species is shown on specimens from eastern Serbia through their internal structures.

Key words: Cretaceous, Late Barremian, eastern Serbia, new species, brachiopods, Kingenidae, Dzirulina.


On the basis of only two specimens found in Upper Barre-
mian sediments of Crnoljevica, the Svrljiške Planine
Mountains, eastern Serbia, Antula (1903) introduced a
new species named Waldheimia  (Zeilleria)  zujovici. Later,
Petković (1930) made the same identification for five
small specimens collected from rocks of the same age at
Bežište on the Suva Planina Mountain. Among the brachi-
opods collected by Petković (1930) from this locality was
a somewhat larger specimen, which he rightly identified as
Waldheimia  (Zeilleria)  pseudojurensis  (Leymerie, 1842).
Note that the above mentioned authors dated the rocks
containing these brachiopods as Hauterivian.

Jankičević (1978, 1979—1980) and Mitrović-Petrović

(1979—1980) later confirmed the Barremian age of the
rocks. More recently, Radulović (2000) and Radulović et
al. (2002) dated them as Late Barremian.

The two specimens described by Antula (1903: p. 47,

pl. 2, figs. 1, 2) are lost and were unavailable for study.
A large display of Lower Cretaceous brachiopods from
a number of east Serbian localities, collected by Zora
Sučić-Protić (1955—1960), has been preserved in the In-
stitute of Regional Geology and Paleontology, Faculty
of Mining and Geology in Belgrade. Another set of
specimens recently collected during detailed geologi-
cal field work in both of the above mentioned localities
was studied. Specimens from Crnoljevica, the same lo-
cality as Antula’s two specimens, include an abundance
of  D.  pseudojurensis, earlier identified by Antula as
Waldheimia  (Zeilleria)  zujovici. Their external and in-
ternal features were studied. Thus it is proposed that the
two specimens described by Antula as Waldheimia
(Zeilleria)  zujovici belong to Dzirulina  pseudojurensis.
The specimens from Bežište, which Petković (1930)
mistakenly believed to be Waldheimia  (Zeilleria)  zujo-

vici, are actually representatives of a new species,
which is described herein.

Geological setting

The brachiopods described in this paper originate from

two localities, Crnoljevica and Bežište, in the Carpatho-
Balkanides of eastern Serbia (Fig. 1). Crnoljevica is locat-
ed in the Kučaj-Svrljig Zone, and Bežište in the
Gornjak-Suva Planina Mountain Zone. Both zones belong
to the Geticum ( = Srednja Gora in Bulgaria).

The sequence at Crnoljevica consists of bioclastic,

marly and argillaceous limestones of Late Barremian

Fig. 1. Location of Upper Barremian brachiopod-bearing sites (aster-
isks) in east Serbian Carpatho-Balkanides. 1 – Crnoljevica, Svrljiške
Planine Mountains; 2 – Bežište, Suva Planina Mountain.

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age. In addition to Dzirulina  pseudojurensis, these
rocks contain many other brachiopod species in associ-
ation with bivalves, echinoids, cephalopods, benthic
foraminifers, and algae (Antula 1903; Radulović 2000;
Radulović & Radulović 2002).

At Bežište, the locality of the new species, the bioclastic

limestones (wackestone and packstone) contain many bra-
chiopods and less abundant bivalves, corals, hydrozoans
and benthic foraminifers. Scleractinian corals and hydro-
zoans from these beds indicate Late Barremian age (Radu-
lović et al. 2002).

The material discussed here is preserved in the first au-

thor’s collection at the Faculty of Mining and Geology,
Belgrade (RGF VR). The synonymy list is presented fol-
lowing the recommendations of Matthews (1973). The su-
perfamily classification follows Williams et al. (1996).

Taxonomic descriptions

Phylum  Brachiopoda Duméril, 1806

Subphylum  Rhynchonelliformea Williams et al., 1996

Class  Rhynchonellata Williams et al., 1996

Order  Terebratulida Waagen, 1883

Suborder  Terebratellidina Muir-Wood, 1955

Superfamily  Kingenoidea Elliott, 1948

Family  Kingenidae Elliott, 1948

Subfamily  Kingeninae Elliott, 1948 [emend. Richardson,


Genus  Dzirulina Nutsubidze, 1945

T y p e   s p e c i e s :

  Terebratula dziroulensis Antula,

1899; Middle Aptian of Georgia.

R e m a r k s : 

On the basis of the Aptian species Terebratu-

la dziroulensis Antula (1899) from western Georgia, Nutsub-
idze (1945) introduced a new genus Dzirulina and showed a
series of transverse sections with spacings from 0.5 to 1 mm,
thus missing important elements of the internal structure,
such as outer and inner hinge plates, crural bases, and hood.

Without insight into the internal structure of Dzirulina,

Smirnova (1960) later described a new genus, Belothyris
from the Early Cretaceous of Beloya, northwestern Cauca-
sus, with B. plana Smirnova (1960) as the type species,
and also two new species of the same genus.

Kvakhadze (1972) used a series of transverse sections

through a specimen of Dzirulina dziroulensis from the
type locality and proved that the genus Belothyris
(Smirnova) was a junior synonym of Dzirulina. This was
also discussed by Owen (1980) and Sandy et al. (1995).

O c c u r r e n c e :

 Hitherto, eleven species of Dzirulina

are known. They have a wide geographical distribution in
the Hauterivian and the Barremian, and a narrower one in
the Aptian, along the northern Tethyan margin (Morocco,
Spain, Sardinia, France, Switzerland, Serbia, western Geor-
gia), and in the Upper Aptian on the southern Tethyan
border (Zululand and probably Pakistan). Beyond the lim-
its of the Tethys, representatives of this genus are known
from the Hauterivian of the Boreal province (southern En-

gland) and from the Lower Albian of the Pacific region
(northern California, Sandy et. al. 1995).

Dzirulina pseudojurensis (Leymerie, 1842)

Figs. 2—5, 6.1—4

*    1842 Terebratula pseudo-jurensis Leymerie, 12, pl. 15, figs. 5, 6
      1843 Terebratula pseudo-jurensis Leym. – Mathéron, 131
.     1847 Terebratula pseudo-jurensis Leym. – d’Orbigny, 74,

pl. 505, figs. 11—16

.    1861 Terebratula  pseudojurensis Leymerie – de Loriol, 121,

pl. 15, figs. 19—21

.    1868 Terebratula  (Waldheimia)  pseudojurensis Leymerie –

de Loriol, 54, pl. 4, figs. 12—14

.   1872 Terebratula  (Waldheimia)  pseudojurensis Leymerie –

Pictet, 93, pl. 203, figs. 11—15

non 1874 Terebratula  (Waldheimia)  pseudojurensis Leymerie –

Davidson, 48, pl. 7, figs. 10—14

.     1903 Waldheimia (Zeilleria) Žujovići Antula, 47, pl. 2, figs. 1, 2
            1907 Zeilleria pseudojurensis Leym. – Karakasch, 216
.v     1930 Waldheimia (Zeilleria) pseudojurensis Leym. – Petković,

107, pl. 3, fig. 1

.v      1953 Waldheimia (Zeilleria) pseudojurensis Antula – Sučić, 110
.v  1953 Terebratula  pseudojurensis Leymerie – Sučić, 111,

pl. 3, figs. 6, 7

.    1970 Belothyris  pseudojurensis (Leymerie) – Owen, 70, pl. 9,

figs. 1—5, 7

.          1975 Belothyris pseudojurensis (Leymerie) – Dieni & Mid-

dlemiss in Dieni, Middlemiss & Owen, 203, pl. 38,
figs. 4—6.

.v   1979—80 Waldheimia  (Zeilleria)  pseudojurensis Leym. –

Jankičević, 225

      1980 Belothyris pseudojurensis (Leymerie) – Owen, 278
.   1989 Zeilleria pseudojurensis (Leymerie) – Gaspard, 84,

pl. 1, fig. 3c

      1990 Belothyris  pseudojurensis (Leymerie) – Smirnova, 134
.    1999 Belothyris  pseudojurensis (Leymerie) – Sulser, 231 with


.    1999 Belothyris  pseudojurensis (Leymerie) – Gaspard, 313,

319, 322, 324, pl. 2, fig. 6

.v   2000 Dzirulina  pseudojurensis (Leymerie) – Radulović, 122,

124, pl. 2, fig. 9

        2003 Belothyris  pseudojurensis (Leymerie) – Gaspard, 266,

fig. 1A

N e o t y p e :

 Since the original specimen described by

Leymerie (1842) is presumed lost, Owen (1980) proposed
another one among the specimens from the Hauterivian
type locality of Marolles (Aube), France.

M a t e r i a l : 

75 complete specimens from Crnoljevica,

the Svrljiške Planine Mountains, 11 specimens from Be-
žište, Suva Planina Mountain.

D e s c r i p t i o n :  External features:

 The exterior of this

species from France was described by Owen (1970). A de-
scription of the morphological characters of specimens
from Serbian Carpatho-Balkanides follows.

The shell is of medium size for the genus, reaching

21 mm in length. In one population, the outline may vary
from elongate-oval, rounded-pentagonal, pentagonal to
almost perfectly circular. The greatest width is situated at
the mid-length of the shell or slightly further anteriorly;
the greatest thickness is in the posterior third. The valves
are nearly equally moderately convex and flattened; the
ventral valve is regularly convex, often with a more or less
developed sulcus on the anterior half; the dorsal valve is

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without a corresponding fold, with a slightly swollen pos-
terior part. The anterior commissure ranges from rectimar-
ginate to slightly uniplicate; the lateral commissure is
gently ventrally arched to straight. The indentation of the
anterior margin is more or less pronounced. The beak is
small, low and wide, erect, in some specimens it is in close
contact with the dorsal umbo, with a relatively small cir-
cular foramen of the mesothyrid to permesothyrid type.
The deltidial plates disjunct. The beak ridges are subangu-
lar and short, border small, with a narrow and concave in-
terarea. The shell surface is marked by well-developed
growth lines, which become more prominent at the mar-

Internal features:

 The internal structure of D. pseudoju-

rensis of the type material has not hitherto been described.
Below is given a detailed description of two specimens
from eastern Serbia (Figs. 2, 3). Recently, Gaspard (2003)
gave transverse serial sections of this species from the Ap-
tian of Auxerre (Yonne, France).

The dental lamellae are ventrally divergent, very short,

nearly reduced before the dorsal valve arises. The hinge
teeth are elongated, deeply inserted in dental sockets. No
cardinal process is developed. The hinge platform is
slightly concave posteriorly, broadening and shallowing
anteriorly, supported by a high septal pillar occupying
about one-half of the valve length, and rapidly decreasing

Fig. 2. Transverse serial sections of Dzirulina pseudojurensis (Leymerie, 1842) through specimen RGF VR 25/114, collected from Upper
Barremian, Crnoljevica, Svrljiške Planine Mountains. Original dimensions of specimens (in mm): L = 16.7, W = 13.2, T = 9.4. Numbers refer
to distance in mm from the ventral apex.

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in height anteriorly of the hood. The platform is formed by
the outer and inner hinge plates, separated by triangular,
dorsally directed crural bases. The crural processes are
ventrally convergent, slightly inwardly concave and mod-
erate in height. The descending branches are much nar-
rower than the slightly ventrally convergent ascending
branches. The transverse band broad, very slightly arched,
with two dorsally directed small processes, joins two as-

cending branches of the loop before it is attached to a septal
pillar. The connecting hood is circular and short, at ap-
proximately one-third of the dorsal valve length. The loop
is kingeniform, thin, extending about three-quarters of the
valve length.

R e m a r k s :

 The species is distinguished externally from

Dzirulina minima sp. nov. by its much larger dimensions,
wide variation in outline, with usually sharp edges. Internal-

Fig. 3. Transverse serial sections of Dzirulina pseudojurensis (Leymerie, 1842) through specimen RGF VR 25/87, collected from Upper
Barremian, Crnoljevica, Svrljiške Planine Mountains. Original dimensions of specimens (in mm): L = 15.5, W = 13.8, T = 9.0. Numbers refer
to distance in mm from the ventral apex.

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ly, it differs in having slightly inclined crural bases, ventrally
convergent crural processes, and wide ascending branches,
and the septal pillar is without a round structure in the axial
part. In five species of genus Dzirulina:  D. pseudojurensis
(Leymerie, 1842), D.  ovula (Anderson, 1938), D. plana
(Smirnova, 1960), D. elliptica (Kvakhadze, 1972), and D.
haughtoni (Owen, 1980), the interiors of which are known,
the septal pillar is without a round structure.

Antula (1903) considered that his new species, Waldhe-

imia  (Zeilleria)  zujovici, which now belong to D. pseudo-
jurensis, to be most similar to Pictet’s (1872: pl. 203,
figs. 11—15) representatives of Waldheimia  (Zeilleria)
pseudojurensis. It differs from the latter in having “a sharp
beak, much greater thickness, protruded anterior end, and
unemarginated lateral edges”. He had only two specimens
(Fig. 4) and could not have known of the highly varied ex-
ternal morphology of this species.

Dieni & Owen (in: Dieni, Middlemiss & Owen 1975) also

described the external morphology of this species from sev-
eral localities of east-central Sardinia. Unlike the typical
form, the specimens mentioned by these authors are much
smaller in size, but are proportionally the same as the origi-
nal specimens of Leymerie (1842). In our opinion, the small
size of the shells is most likely the result of unfavourable
living conditions. They also wrote that specimens of this
species collected from different localities of the Swiss and
French Jura could vary widely in size, from small, as the Sar-
dinian specimens, to large, as the type forms. For these au-
thors, the shell size was not of specific importance.

We also found that the forms of this species from differ-

ent localities of eastern Serbia vary greatly in size, and in
outline. In a study of the external morphology on a large
number of specimens from eastern Serbia, the following
was noted: – the outline varies greatly, from elongate oval,
rounded pentagonal, pentagonal to circular (Fig. 5); – the
sulcus is on the anterior half of the shell, and the indenta-
tion in the anterior margin is more or less marked; – the
shell size varies from small to large; – the valves are flat-
tened, moderately or strongly convex; – very rarely some
marginal thickening develops in adult specimens.

O c c u r r e n c e :

 The species has been recorded from the

Hauterivian of Morocco, eastern Iberides, east-central Sar-
dinia, the South Paris Basin, the French and Swiss Jura,
Valanginian and Hauterivian of the Crimea and Caucasus,
Late Barremian of eastern Serbia (Fig. 5).

Dzirulina minima sp. nov.

Figs. 6.5—10, 7

non   1903 Waldheimia (Zeilleria) Žujovići Antula, 47, pl. 2, figs. 1, 2

(= Dzirulina  pseudojurensis Leymerie)

.v        1930 Waldheimia (Zeilleria) Žujovići Antula – Petković, 107,

pl. 3, fig. 4

non v  1953 Waldheimia (Zeilleria) žujovići Antula – Sučić, 110

(= Dzirulina  pseudojurensis Leymerie)

non v  1978 Waldheimia  žujovići Antula – Jankičević, 129 (= Dzi-

rulina  pseudojurensis Leymerie)

D e r i v a t i o   n o m i n i s : 

Latin word “minima” – small,

referring to the shell size.

H o l o t y p e :

 Specimen RGF VR 76/8, figured in

Fig. 6.7. Dimensions (in mm): length = 10.8,  width = 9.3,
thickness = 6.8.

P a r a t y p e s :

 The specimens RGF VR 76/6, 7, 9, 10, 11,

figured in Figs. 6.5, 6, 8—10.

S t r a t u m   t y p i c u m : 

Late Barremian bioclastic lime-


L o c u s   t y p i c u s :

 Bežište, Suva Planina Mountain,

eastern Serbia.

D i a g n o s i s :

 Small Dzirulina species with a pentagonal

outline, crowded growth lines along a marginal shell
thickening, round structure in the axial part of the septal
pillar, strongly dorsally inclined crural bases, and subpar-
allel crural processes.

D e s c r i p t i o n :  External features:

 Shell small in size

for the genus, largest specimen 12.1 mm in length,
11.2 mm in width, 5.7 mm in thickness, always longer
than wide, distinctly pentagonal in outline, truncated len-

Fig. 4. Antula’s (1903: pl. 2, figs. 1, 2) original drawings of two
specimens of Dzirulina pseudojurensis (Leymerie,  1842),  earlier re-
ferred to by Antula as Waldheimia  (Zeilleria)  zujovici, from Upper
Barremian, Crnoljevica, Svrljške Planine Mountains. a—d – young
adult specimen, e—h – fully adult specimen. Both specimens are il-
lustrated in dorsal, ventral, lateral and anterior views and natural size.

Fig. 5.  Shape variability in Dzirulina pseudojurensis (Leymerie, 1842) of specimens, collected from Upper Barremian, Crnoljevica, Svrl-
jiške Planine Mountains. Scale bar = 10 mm.

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Fig. 6. 1—4 – Dzirulina pseudojurensis (Leymerie, 1842). 1 – RGF VR 76/4, Late Barremian, Bežište, Suva Planina Mountain; length of
shell 16.3 mm. 2 – RGF VR 25/147, Late Barremian, Crnoljevica, Svrljiške Planine Mountains; length of shell 16.5 mm. 3 – RGF VR
25/137 Late Barremian, Crnoljevica, Svrljiške Planine Mountains; length of shell 17.2 mm. 4 – RGF VR 76/5, Late Barremian, Bežište,
Suva Planina Mountain; length of shell 21.0 mm. 5—10 – Dzirulina minima sp. nov., Late Barremian, Bežište, Suva Planina Mountain.
5 – RGF VR 76/6, paratype; length of shell 9.6 mm. 6 – RGF VR 76/7, paratype, used for transverse serial sections; length of shell
10.6 mm. 7 – RGF VR 76/8, holotype; length of shell 10.8 mm. 8 – RGF VR 76/9, paratype; length of shell 11.0 mm. 9 – RGF VR 76/10,
paratype; length of shell 11.1 mm. 10 – RGF VR 76/11, paratype; length of shell 12.1 mm. All figured specimens were coated with am-
monium chloride before being photographed. The letters imply the view: a – dorsal, b – ventral, c – lateral, d – anterior. Photo-
graphs by V. Radulović.

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Fig. 7. Transverse serial sections of Dzirulina minima sp. nov. through specimen RGF VR 76/7, collected from Upper Barremian, Bežište,
Suva Planina Mountain. Original dimensions of specimens (in mm): L = 10.6, W = 9.8, T = 6.1. Figured on Fig. 6.6. Numbers refer to distance
in mm from the ventral apex. Enlarged hinge plates, crural bases, and round structure (arrow) in the axial part of the septal pillar are shown
above the original structures


in section 2.5. Due to the small size of the shell, the transverse band most probably invisible during sectioning.

ticular in the anterior contour, anterior end narrower and
indentate. Both valves are moderately, and equally bicon-
vex. Greatest width and thickness at the mid-length of the
shell. Beak moderately produced, erect to slightly in-
curved, in close contact with the dorsal umbo, with short
and rather distinct beak ridges, foramen circular, mesothy-
rid. Interarea in some specimens relatively large, concave.
Each valve has two low, barely marked short ridges sepa-
rated by a more or less developed sulcus on the anterior
half. The anterior commissure is rectimarginate to incipi-
ently uniplicate; the lateral commissures are straight or

slightly ventrally arched. Marginal thickening develops
in the gerontic stage. Numerous fine growth lines concen-
trated towards the anterior and lateral commissures. Entire
surface finely punctated.

Internal features: 

Two specimens were sectioned. The

first specimen revealed a broken loop. The second speci-
men with a complete loop is figured (Fig. 7).

The dental lamellae are thin and short, ventrally diver-

gent, supporting elongated hinge teeth, which become
bracket-like, in cross-section, anteriorly. The sockets are
short, but deep. The outer hinge plates are not clearly de-

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limited from the inner socket ridges. The inner hinge
plates are thin, conjunct, slightly dorsally inclined. They
are separated by triangular in cross-section, strongly dor-
sally inclined crural bases; together forming a broad, con-
cave and shallow hinge platform. The platform is
supported by a thin, long, blade-like, septal pillar, reach-
ing about mid-length of the valve, highest at the level
where it is linked with the ascending branches. Posterior-
ly, the septal pillar possess a round structure in the axial
part. The crural processes are subparallel, relatively low.
The proximal ends of the ascending branches widening,
attached to the septal pillar, building a very short and cir-
cular hood. The distal ends of the descending and ascend-
ing branches are of the same thickness. The loop is long,
about three-quarters of the dorsal valve length.

R e m a r k s :

 Among the species of Dzirulina, the new

species can only be compared with D. pseudojurensis from
which it differs in being much smaller, always pentagonal
in outline, in having dense and constantly strong growth
lines along the lateral and anterior commissures, and mar-
ginal shell thickening. On the other hand, it can be distin-
guished internally from the latter by its round structure in
the axial part of the septal pillar, much more inclined cru-
ral bases, subparallel crural processes, and a loop consist-
ing of descending and ascending branches of the same
thickness. The main external features which connect both
species are a more or less developed sulcus on the anterior
half and an indentate anterior margin.

All six specimens of the new species possessed small

shells. Though small, the shell bears several prominent
growth lines on its anterior end, which is, without any
doubt, a feature of the adult stage. Hence, the inference is
that the forms of the new species are much smaller than
other representatives of the same genus. Relatively large
specimens of Dzirulina pseudojurensis (Fig. 6.4) were
found in the same layers of the Bežište locality, from
where the small specimens of the new species were recov-
ered. Therefore, the small shells of these new species can-
not be attributed to the prevailing ecological conditions
and it is clearly a morphogenetic feature.

Radulović (1991) described specimens of three species:

Agerithyris minima,  Minithyris  rhomboidalis, and Struve-
thyris  elongata from the Middle Jurassic of the Laz locali-
ty, eastern Serbia, which also had densely-set growth lines
near the anterior commissure and possessed small shells,
and confirmed they have a gerontic aspect. The size of
these adult small shells, found in association with other
rhynchonellides and terebratulides of normal size, cannot
be attributed to detrimental life conditions.


The Upper Barremian shallow-water limestones of the

eastern Serbian Carpatho-Balkanides contain abundant
brachiopods. The commonest among them is Dzirulina
pseudojurensis, which has a wide geographical distribu-
tion in the Hauterivian and Barremian along the northern
Tethyan margin.

The interior features and variation of  the external mor-

phology (size, outline, valve convexity, sulcus and inden-
tation development) of the species are described using a
large number of specimens from many localities of eastern

Another representative of this genus, Dziruline minima

sp. n., was identified earlier by Antula (1903) as Waldheim-
ia  (Zeilleria)  pseudojurensis.  The new species  distinctly
differs from any other Dzirulina species in its small shell,
dense and strong growth lines along flattened shell edges,
and round structure in the axial part of the septal pillar. It is
known only from the locality of Bežište, eastern Serbia.


We wish to thank Dr. Eric Simon

(Bruxelles) for his helpful and valuable suggestions for
improvements. We are grateful to the reviewers: Prof. Dr.
Daniele Gaspard (Paris), Dr. Jozef Michalík (Bratislava)
and Dr. Attila Vörös (Budapest) for their constructive com-
ments on the manuscript. Msc. Lynne Katsikas (Belgrade)
corrected the English. The research has been supported by
Ministry of Science and Environmental Protection of the
Republic of Serbia, Project No. 146023.


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