GEOLOGICA CARPATHICA, OCTOBER 2010, 61, 5, 355—364 doi: 10.2478/v10096-010-0021-4
Paleozoic ammonoid faunas are known from only a few places
in the Balkan Peninsula. The longest known of these is the oc-
currence of early late Visean ammonoids at Prača near Saraje-
vo, from which Kittl (1904a) described the peculiar Entogonites
grimmeri. This strange ammonoid, which possesses tetrangu-
larly coiled inner whorls, has been subsequently discovered in
many places (e.g. Rhenish Mountains, British Isles, Anti-Atlas
of Morocco, Alaska, Great Basin of Utah) and serves as an im-
portant index species for the intercontinental correlation of
early late Visean (Mississippian) sediments.
Entogonites was also collected at Milivojevića Kamenjar
site in Družetić village near Valjevo, but these records were
misinterpreted by J. Kullmann in Stevanović & Kullmann
(1962), who considered these specimens to be homeomor-
phic and therefore not related to Entogonites. Instead, these
fossils were erroneously attributed to the late Bashkirian ge-
nus Gastrioceras, for which J. Kullmann (op. cit., p. 88) in-
troduced the new subgenus Branneroceratoides.
The outcrop in the Družetić area (Fig. 1) is important for a
number of reasons. It is the richest Carboniferous ammonoid
locality in the Balkan Peninsula, situated in a region, which is
not fully understood in terms of its paleogeographic position
between Laurussia and Gondwana. It contains two productive
ammonoid horizons, allowing precise biostratigraphic assign-
ment and thus a correlation with time equivalent faunas from
Northern and Western Europe, North Africa, the Urals,
Novaya Zemlya, etc. The succession is fully composed of
limestones, enabling the sampling of conodonts and thus the
chance to study a second fossil group for biostratigraphy.
In a series of papers the results of a re-study of the am-
monoid and conodont faunas from the Milivojevića Kamenjar
Early late Visean ammonoid faunas from the Jadar Block
, DIVNA JOVANOVIĆ
, MATEVŽ NOVAK
and MILAN N. SUDAR
Museum für Naturkunde, Leibniz Institute at the Humboldt University Berlin, Invalidenstraße 43, 10115 Berlin, Germany;
Geological Institute of Serbia, Rovinjska 12, 11000 Belgrade, Serbia; email@example.com
Geological Survey of Slovenia, Dimičeva 14, SI-1000 Ljubljana, Slovenia; firstname.lastname@example.org
Department of Paleontology, Faculty of Mining and Geology, University of Belgrade, Kamenička 6, PO Box 62, 11000 Belgrade, Serbia;
(Manuscript received February 5, 2010; accepted in June 10, 2010)
Abstract: The outcrop at Milivojevića Kamenjar in Družetić (Jadar Block, Vardar Zone, NW Serbia), which exposes a
fossiliferous limestone olistolith, is one of the key sites for Carboniferous stratigraphy and paleogeography in the
Balkan Peninsula. Its age has been debated several times, and re-examination of the succession was required. Based on
ammonoids and conodonts, an interval spanning from the latest Devonian to the basal Serpukhovian is represented.
From the early late Visean portion of the section, the new ammonoid genus and species Ubites filipovici gen. nov. et sp.
nov. is described. Entogonites tetragonus (Kullmann, 1962), a formerly misinterpreted ammonoid species, is revised.
Key words: Mississippian, Visean, Jadar Block, Vardar Zone, NW Serbia, biostratigraphy, taxonomy, Ammonoidea.
section and age determination, based on the current state of
knowledge, will be presented (e.g. Korn et al. in print). In
this first paper some of the newly determinated ammonoids
from the “upper ammonoid horizon” will be described, and
also the new stratigraphic position of this important fossil
horizon will be established.
Geographical position, geological context, and
general stratigraphy of the outcrop in Družetić
The Paleozoic sedimentary rocks exposed in the area of
Družetić village belong to the Jadar Block, which is geo-
graphically located on the southern margin of the Pannonian
Basin: mostly in north-western Serbia, southern Srem, and
partially westward over the Drina River in eastern Bosnia
(Fig. 1). The name of this tectonostratigraphic unit, which is
now a part of the Vardar Zone, derived from the “Jadar de-
velopment of Paleozoic” (Simić 1938).
The Jadar Block is an isolated, exotic block terrane, where
Dinaridic features predominate. It was incorporated into the
Vardar Zone before the Late Cretaceous (Karamata et al.
2000; Karamata 2006, etc.). It is surrounded by the Vardar
Zone Western Belt, except for the south-easternmost part,
where it is in direct contact with the Kopaonik Block and
Ridge Unit, which is also a part of the Vardar Zone (Fig. 1). In
this area, deposition of sediments took place during the
Variscan and early Alpine evolution with obvious similarities
to time-equivalent successions of the “Bükkium” (NE Hunga-
ry), the Sana-Una terranes (NW Bosnia and Herzegovina), and
even the Carnic Alps (Protić et al. 2000; Filipović et al. 2003).
Carboniferous formations of the Jadar Block, as part of the
late Variscan sedimentary cycles, comprise both autochtho-
KORN, JOVANOVIĆ, NOVAK and SUDAR
nous and allochthonous marine rocks. The former, devel-
oped in the Jadar Autochthon (Krupanj-Valjevo, Vlašić, Slo-
vac, and Ub Units), are divided by structural and facial
criteria into clastic flysch (Variscan flysch or Vlašić Forma-
tion), pelagic carbonates (Družetić Formation), and molasse
sequences (Ivovik Formation, Kriva Reka Formation)
(Fig. 2). The rocks of the Jadar Allochthon (Likodra Nappe)
are characterized by an apparently continuous succession of
deep-water deposits (Variscan flysch) followed by transi-
tional, basinal carbonates (Đulim Formation) and shallow
marine carbonate-terrigenous sediments mainly with bioher-
mal characteristics (Rudine Formation, Stojkovići Forma-
tion, and Stolice Limestone Formation) (Fig. 2).
One of the main characteristics of the late Variscan
succession in the Jadar Block is molasse-type sediments,
which are deposited only in the Jadar Autochthon (Filipović
1995; Filipović et al. 2003). The terrestrial debris flow-type
sediments of the Ivovik Formation (Krupanj-Valjevo, Vlašić
and Ub Units), were formed first and the marine offshore
thick fusulinacean carbonates without siliciclastics of the
Kriva Reka Formation (only in the Krupanj-Valjevo Unit on the
Jadar Paleozoic southern margin) were deposited later. The
Ivovik Formation was deposited in the middle parts of the
Moscovian (Podolskian). The age of the Kriva Reka Formation
is from late Moscovian (Myachkovian) to Early Permian
(Asselian). It lies between the olistostrome Ivovik Formation
and the transgressive Middle Permian clastics (Bobova Breccia
and Cerova Formation) (Filipović et al. 2003) (Fig. 2).
The Ub Unit, as the north-eastern extension of the Jadar
Autochthon, occupies a large area on the left side of the Ub
River in the Tamnava Basin, where it is mostly covered with
Neogene and Triassic sediments. During the Middle Devonian
to Early Carboniferous (Mississippian), pelagic limestones of
the Družetić Formation were deposited on the intrabasinal
Fig. 1. Location of the Milivojevića Kamenjar section, Jadar Block, Vardar Zone (NW Serbia). A – Terranes of part of the Balkan Penin-
sula (Karamata et al. 2000): SMCT – Serbian-Macedonian Composite Terrane; MVZ – Main Vardar Zone; KBRU – Kopaonik Block
and Ridge Unit; VZWB – Vardar Zone Western Belt; JBT – Jadar Block Terrane; DIE – Drina-Ivanjica Element; DOB – Dinaridic
Ophiolite Belt; EBDT – East Bosnian-Durmitor Terrane. B – Units of the Jadar Block Terrane (simplified from Filipović et al. 2003):
KV – Krupanj-Valjevo; V – Vlašić; S – Slovac; U – Ub; LN – Likodra Nappe.
EARLY LATE VISEAN AMMONOID FAUNAS (NW SERBIA)
rise. Subsequently, mountain massifs of the Ub Unit were
formed in connection with the Asturian orogeny. They follow
transgressively over the mentioned pelagic limestones as the
products of gravity sliding (siltstones with limestone clasts
and olistoliths) followed by alternations of massive, bedded
and thin-bedded silty limestones with woody plant remains.
These deposits were named the Ivovik Formation according to
their analogous developments with the same Formation in the
Krupanj-Valjevo Unit (Filipović 1995). In the Ub Unit the
deposition finished with these sediments.
In the wider area of the village of Družetić, 24 km north of
Valjevo near the road Valjevo-Koceljeva-Šabac and on the
north-eastern slopes of the Vlašić Mountains (NW Serbia)
(Fig. 1), the older, olistostromal part of the Ivovik Formation
is developed. It is made up of a siltstone matrix with clasts
and olistoliths of Devonian and Mississippian limestones of
the Družetić Formation. In previous investigations, Devo-
nian limestones were found at many places in NW Serbia
(Filipović et al. 1975), and even in Družetić, they were treat-
ed as autochthonous. According to recent data they are in
fact olistoliths and belong to the younger Ivovik Formation
The best data of the latest Devonian and Mississippian
parts of the Družetić Formation are derived from the inverse
olistolith in the Milivojevića Kamenjar section, which is
located on the right side of the Ub River in the south-western
area of Družetić village (coordinates: x 4920
44.4191 N, 19.8128 E; Fig. 3). It is known as the only
undisputed locality of a Namurian goniatite fauna in the
Balkan Peninsula, where Stevanović & Kullmann (1962)
attributed the cephalopod faunas from two fossiliferous
horizons to the “upper Eumorphoceras Stufe” (E
) within the
older part of Namurian A. A record of Dombarites, determined
by Kullmann and mentioned in Veselinović & Filipović (1989)
Fig. 2. Carboniferous formations developed in the Jadar Block during the Variscan (only Carboniferous part) and late Variscan sedimentary cycles
(modified and simplified from Filipović et al. 2003). Abbreviations and legend: SS – Stupnica Sandstone Formation; ŽF – Županjac Fm;
1a – bathyal, pelagic carbonates (Družetić Fm); 1b – flysch (turbidites), flysch-like sediments (Variscan Flysch, Vlašić Fm, Stupnica Sandstone
Fm, Županjac Fm); 2 – open shelf, basinal, platform slope carbonates (Đulim Fm); 3 – shallow marine carbonates (Rudine Fm, Stolice Lime-
stone Fm); 4 – intertidal and shallow subtidal clastics (Stojkovići Fm); 5 – molasse-type sediments; 5a – (Ivovik Fm), 5b – (Kriva Reka Fm).
KORN, JOVANOVIĆ, NOVAK and SUDAR
as well as Djordjijevski-Kalambokis et al. (1990) may be
Serpukhovian in age, but the specimen could not be traced.
Later, Ruzhencev & Bogoslovskaya (1971) noted some
mistakes within the original determinations (of some species
and genera), which made a revision of the ammonoid fauna
and of the whole section necessary.
The rock succession of the Milivojevića Kamenjar
At Milivojevića Kamenjar in Družetić, approximately 15
meters of sedimentary rocks, almost exclusively carbonates,
are exposed. The bedding planes dip 10 to 25° in a south-
western direction. The succession can be subdivided into
five units, in descending order (i.e. stratigraphically from
older to younger) in the outcrop (Fig. 3):
1. Dark grey, nodular limestones (> 2 m) with a high clay
content, in part only poorly exposed. The limestone nodules
and their macrofossil content are strongly deformed.
2. Grey bedded, also nodular lime-
stones with a minor clay content
3. Partly fossil-rich dark grey bedded
limestones (0.65 m), which possess a
high dolomite content. Two fossil sam-
ples (DRZ 1 and DRZ 2 separated by a
distance of about 20 cm) were taken
from the “upper” (i.e. stratigraphically
older) portion of the unit. Particularly
sample DRZ 2 is a coquina of a mass
occurrence of ammonoids, but most of
the specimens larger than 10 mm are
4. Light grey, thick-bedded or almost
non-bedded non-fossiliferous micritic
limestones (8.60 m).
5. Grey, well-bedded micritic nodular
limestones (2.80 m) with a minor clay
content. In this unit, five ammonoid sam-
ples from the middle 1.50 m were ob-
tained. The occurrence of the macrofossils
is patchy, and in some cases coquinas
packed with ammonoids occur in more
sparitic portions of the section.
The stratigraphy of the
Milivojevića Kamenjar section
The discussion about the stratigraphic
age of the fossiliferous limestone occur-
rences at the Milivojevića Kamenjar site
in Družetić began parallel to the de-
scription of the outcrop. On the basis of
the two ammonoid faunas, Stevanović
& Kullmann (1962) postulated that the
entire outcrop belongs in the Namurian
(“obere Eumorphoceras Stufe”). Doubts
Fig. 3. Milivojevića Kamenjar section (modified from Filipović 1995, Filipović et al. 2008,
and incorporating the new data). A – Sketch of the geographical position in the area of
Družetić village (Jadar Block, Vardar Zone, NW Serbia). B – Section of the Milivojevića
Kamenjar olistolith. Legend: Unit 1 – Dark grey nodular limestones with thin shaly beds
(Famennian according to Filipović 1995); Unit 2 – Grey bedded nodular limestones (Tour-
naisian according to Filipović 1995); Unit 3 – Grey bedded limestones with ammonoids
(Namurian “upper fossiliferous layer” according to Stevanović & Kullmann 1962; early late
Visean according to our investigation); Unit 4 – Grey massive, thick-bedded and bedded
limestones (Visean according to Filipović 1995); Unit 5 – Grey bedded limestones with am-
monoids (Namurian “lower fossiliferous layer” according to Stevanović & Kullmann 1962;
Lower Serpukhovian according to Filipović 1995; latest Visean to earliest Serpukhovian in
this paper); 6 – Podolskian olistostromal deposits of the Ivovik Formation (Filipović 1995).
C – Detail of the upper part of unit 3 (“upper ammonoid horizon”, Entogonites Genus Zone
of the early late Visean (middle Asbian) in this paper).
were cast by I. Filipović, who found Late Devonian microfos-
sils (styliolinids, ostracods) in the nodular limestones such as
the horizons immediately “above” the massive limestones of
the section. These arguments were dismissed by Stevanović &
Kullmann (1962, footnotes on p. 50 and 66), who insisted on a
Namurian age of the beds under discussion.
Subsequent investigations and studies of conodonts from
the section undermined the original statement that only
Namurian sedimentary rocks exist in the limestones of the
Milivojevića Kamenjar section. First, Spasov & Filipović
(1967) found besides Namurian conodonts also Late Devonian
(Famennian) conodonts above the “upper fossiliferous
layer”. Detailed, but rather different results were presented by
Scharfe (1977, p. 26), who found Famennian, Tournaisian,
and Visean conodonts in the same section; he stated
“Aufgrund von Conodonten konnte das Namur in Družetic
aber bisher nicht nachgewiesen werden”.
After this, detailed stratigraphic results from the
Milivojevića Kamenjar site were realized by investigations
of conodonts (Stojanović-Kuzenko & Pajić in Filipović
EARLY LATE VISEAN AMMONOID FAUNAS (NW SERBIA)
1995). Following olistrostromal deposits of the Ivovik Forma-
tion, the “uppermost” parts of the limestone olistolith of 16 m
thickness consist of latest Famennian nodular limestones with
conodonts from lower-middle parts of the Siphonodella
praesulcata Zone. Downward are Tournaisian bedded lime-
stones, from which Siphonodella sulcata, S. duplicata and S.
sandbergi zone conodonts were determined. The latest Tour-
naisian conodont zone with Scaliognathus anchoralis was
found in the following parts of the grey bedded limestones
near the second, “upper fossiliferous layer”, which is, accord-
ing to Stevanović & Kullmann (1962), Namurian in age. Be-
low it, the next, largest part of the olistolith (thickness 12 m) is
made up of massive, thick-bedded and bedded limestones
where the next conodont zones of Visean age were deter-
mined: Gnathodus texanus, Gnathodus bilineatus, and
Lochriea nodosa. The final, “lowermost” parts of the section,
i.e. the “lower fossiliferous layer” of Stevanović & Kullmann
(1962), contains conodonts of the Kladognathus—Gnathodus
girtyi group Zone. It belongs to the lower part of the Ser-
pukhovian (Stojanović-Kuzenko & Pajić in Filipović 1995).
At this moment new and more detailed investigations of
the conodonts are in progress.
The ammonoid faunas from Milivojevića Kamenjar
Carboniferous ammonoids from the Milivojevića Kamenjar
section in Družetić were first discovered in 1956 by P.
Stevanović, who called the beds with these fossils cephalo-
pod limestone (Stevanović 1962). J. Kullmann then studied
the ammonoids; and then together with Stevanović pub-
lished an extensive monograph paper (Stevanović &
Kullmann 1962) and interpreted the faunas according to their
stratigraphic age. According to this study, two ammonoid
horizons were distinguished (Fig. 4):
A “lower horizon” (which is in fact the stratigraphically
younger, uppermost Visean to lower Serpukhovian horizon)
was characterized by the following species:
“Prionoceras (Irinoceras) stevanovici”,
“Proshumardites (Proshumardites) serbicus”,
“Proshumardites (Trigonoshumardites) wocklumerioides”,
“Pronorites uralensis uralensis”.
An “upper horizon” (which is in fact the older, lower up-
per Visean horizon) yielded the following species:
“Gastrioceras (Branneroceras) branneri branneri”.
Finally, five species were reported to occur in both horizons:
“Cravenoceras arcticum subinvolutum”,
“Gastrioceras (Branneroceratoides) tetragonum”,
This last category requires discussion. According to the
new investigation of the Milivojevića Kamenjar site, not a
single species was found to occur in both horizons. The sig-
nificant stratigraphic difference between the two assemblages,
which was not recognized by Stevanović & Kullmann
(1962), makes it very unlikely that there are species, which
are present in both of them. It appears that in some cases, as
in “Gastrioceras (Branneroceratoides) tetragonum”, mixing
of samples in the outcrop caused the erroneous record of this
species in the “lower horizon”. In other cases, misinterpreta-
tions may be responsible. The species putatively occurring in
both horizons were possibly the reason why J. Kullmann (in
Stevanović & Kullmann 1962, p. 65) concluded that “the
difference between the two beds, according to their faunas, is
insignificant” (“… Unterschied zwischen diesen beiden
Schichten in faunistischer Hinsicht unbedeutend.”). It is pos-
sibly the reason why J. Kullmann did not recognise the
Visean age of the “upper horizon”.
In the faunal list of Stevanović & Kullmann (1962) it is visi-
ble that names of species from very different regions have
been used, the South Urals (3 species), Novaya Zemlya (1),
Northern England (1), and the American Midcontinent (1). A
further six species have been newly described. On the basis
of this, J. Kullmann concluded that the closest relationships
are noticeable with the Urals (op. cit., p. 68), but also stated
that the fauna from Družetić has an intermediate position be-
tween the “epicontinental and geosynclinal facies” of the
Carboniferous occurrences in Europe.
Newly collected material
Two field sessions were carried out to resample the
Milivojevića Kamenjar section in Družetić. The aim of the
fieldwork was to obtain precise bed-by-bed collected material
of the two fossil horizons that were described by Stevanović
& Kullmann (1962). Both horizons, which are separated by
9.70 meters of rocks largely without macrofossils, were rec-
ognized in the section and could even be subdivided, with
the “lower horizon” providing five successive samples within
a range of 1.50 m (samples DRZ A to DRZ E in stratigraphi-
cally ascending order), and the “upper horizon” separated in
two units 20 cm apart (with sample DRZ 2 being the strati-
graphically older horizon and sample DRZ 1 the younger;
The two samples DRZ 1 and DRZ 2 differ slightly in their
ammonoid spectrum. Entogonites and Ubites gen. nov. occur
in both, but with significantly different numbers. The older
sample DRZ 2 contains Ubites filipovici gen. nov. et sp. nov.
as the predominant species, whereas Entogonites is very
rare. Sample DRZ-1 yielded Ubites gen. nov. in very low
numbers, but Entogonites is very abundant. Apart from the
species E. tetragonum, which is revised here, two other spe-
cies of the genus occur. These will be described in a subse-
A very precise age determination can be made on the basis of
the occurrence of Entogonites. The genus is particularly known
from the Rhenish Mountains of Germany, where its stratigraph-
ic occurrence is best documented (Nicolaus 1963; Korn 1988;
Korn & Horn 1997). The two successive species E. nasutus
and E. grimmeri are index species of two successive biozones in
the early Asbian (early late Visean). E. grimmeri shows a co-oc-
currence with the oldest species of Goniatites, G. hudsoni Bisat,
1934 in only one single nodular horizon.
KORN, JOVANOVIĆ, NOVAK and SUDAR
The North African species Entogonites saharensis can be
found in the body chambers of the species Goniatites lazarus
Korn, Klug & Mapes, 2005, together with species of the gen-
era Maxigoniatites, Bollandoceras, and Calygirtyoceras
(Korn et al. 2007), which form a characteristic assemblage of
the early late Visean.
Occurrences of Entogonites in Alaska and Utah (Gordon
1957) largely confirm these results. E. borealis occurs in the
two regions with Goniatites americanus Gordon, 1971. Tak-
ing all this data into account, there is no doubt that samples
DRZ 1 and DRZ 2 from Družetić possess an early late
The main results of the new investigations of the outcrop
are (Fig. 4):
(1) The “lower” and the “upper horizon” contain completely
different ammonoid species and genera; there is not a single
species, which occurs in both horizons. It can even be stated
that the faunas are so different from one another that a confu-
sion of the two is impossible even in surface collected material.
(2) The “lower” and the “upper horizon” are lithologically
markedly different, the “lower horizon” is a dense and fine-
grained light grey nodular limestone with occasional fossil
content and the “upper horizon” is a coarse-grained dolomitic
limestone packed with ammonoids in two clearly separable
coquinas. These features also make it very easy to separate
the two horizons.
(3) The “lower horizon” is the stratigraphically younger of
the two horizons.
(4) The “upper horizon” is early late Visean (middle Asbian)
in age, the “lower horizon” belongs to latest Visean (upper-
most Brigatian) and the early Serpukhovian (Pendleian).
(5) The “lower horizon” (latest Visean to early Ser-
pukhovian) shows close relationships to time equivalents of
the Cantabrian Mountains and South Urals.
(6) Sample DRZ 2 (the stratigraphically older) of the “up-
per horizon” contains a fauna composed of Ubites gen. nov.
(which is the most abundant genus in this sample), very rare
Entogonites, Beyrichoceras, Bollandites(?), and yet undeter-
mined prolecanitid ammonoids.
(7) Sample DRZ 1 contains very abundant specimens of
Entogonites (E. tetragonum as well as one or two additional
species of the genus) and prolecanitids. Ubites gen. nov. is
rare in this sample.
(8) Samples DRZ A to DRZ E contain, among others, the
ammonoid genera Dombarites and Ophilyroceras as well as
prolecanitid and pronoritid species.
Systematic paleontology (D. Korn, M. Sudar)
Order: Goniatitida de Haan, 1825
Superfamily: Girtyocerataceae Wedekind, 1918
Family: Entogonitidae Ruzhencev & Bogoslovskaya, 1971
F a m i l y d e f i n i t i o n : Girtyocerataceae with strong
radial ornament consisting of sharp ribs, which frequently
Fig. 4. Visean and partly Serpukhovian chronostratigraphy, estimated geochronological scale (after Korn & Kaufmann 2009), conodont
succession, ammonoid zonation for the Rhenish Mountains and their duration, global and South Urals ammonoid genus zones (after
Ruzhencev & Bogoslovskaya 1971; Korn et al. 2007), and position of the ammonoid-bearing horizons in the Milivojevića Kamenjar olis-
tolith. SE. – Serpukhovian; H. – Holkerian; P. – Pendleian.
EARLY LATE VISEAN AMMONOID FAUNAS (NW SERBIA)
dichotomize and form a ventrolateral projection and a deep
ventral sinus. Inner whorls tetrangularly coiled in many
Entogonites Kittl, 1904b
Tetragonites Kittl, 1904a [homonym of Tetragonites
Kittliella Frech, 1906 [synonym of Entogonites Kittl, 1904]
Ubites gen. nov.
D i s c u s s i o n : Entogonites is still an enigmatic genus in
the Early Carboniferous ammonoid faunas, and its phyloge-
netic relationships are not resolved. Traditionally, the genus
has been placed near Nomismoceras, based on the thinly dis-
coidal, relatively widely umbilicate conch, the ornament
with rather high ventrolateral projection, and the suture line
with rounded adventive lobe.
With the discovery of the new genus Ubites, Entogonites
can be connected phylogenetically with other Early Carbonif-
erous ammonoids, suggesting that Nomismoceras is probably
not a close relative. Ubites gen. nov., which possesses an or-
nament like Entogonites but an ontogeny with an evolute
ribbed juvenile stage and closure of the umbilicus in the later
ontogenetic stage (similar to Calygirtyoceras), may link the
family Entogonitidae with ammonoids such as the Girtyocera-
tidae. The family Entogonitidae is therefore tentatively placed
in the superfamily Girtyocerataceae.
Genus: Entogonites Kittl, 1904
T y p e s p e c i e s : Tetragonites Grimmeri Kittl, 1904a
G e n u s d e f i n i t i o n : Entogonitidae with simple ontoge-
ny. Adult whorls slightly narrower umbilicate; inner whorls
borealis: Entogonites borealis Gordon, 1957, p. 53; Alaska
grimmeri: Tetragonites Grimmeri Kittl, 1904, p. 677; Bosnia
nasutus: Pericyclus nasutus Schmidt, 1941, p. 151; Harz
saharensis: Entogonites saharensis Korn, Klug & Mapes,
2005, p. 363; Anti-Atlas
tetragonus: Gastrioceras (Branneroceratoides) tetragonum
Kullmann, 1962, p. 88; NW Serbia.
Entogonites tetragonus (Kullmann, 1962)
1962 Gastrioceras (Branneroceratoides) tetragonum n. sp. Kullmann
in Stevanović & Kullmann, p. 88—90, pl. 2, figs. 2—6
H o l o t y p e : Specimen SLM 4063/38 (Natural History
Museum Belgrade), illustrated by Stevanović & Kullmann
(1962) on pl. 2, fig. 2.
T y p e l o c a l i t y a n d h o r i z o n : Milivojevića Kamenjar
site in Družetić (NW Serbia); “upper horizon”, Entogonites
M a t e r i a l : Thirty paratypes in the Natural History Muse-
um Belgrade and 25 newly collected specimens; conch di-
ameter up to 9.5 mm.
D i a g n o s i s : Species of Entogonites with a thinly discoidal
conch (ww/dm = 0.35—0.40), moderately wide to wide umbili-
cus (uw/dm = 0.40—0.50) and broadly rounded venter at 8 mm
conch diameter. Aperture moderately low (WER = 1.80—1.90).
Inner whorls with moderate tetrangular coiling. Shell orna-
ment with 35 moderately strong, rounded and ventrolaterally
dichotomizing ribs. Ventrolateral projection of ribs low, ven-
tral sinus moderately deep.
D e s c r i p t i o n : Specimen MB.C.19128.1 is a rather well-
preserved specimen with 8.5 mm conch diameter. It is thinly
discoidal and evolute (ww/dm = 0.36; uw/dm = 0.46) and pos-
sesses a moderately high aperture (WER = 1.83). The inner
whorls are inconspicuously tetrangularly coiled. About 30
rounded ribs can be counted on the last volution; they extend
in a linear course across the inner flank and bifurcate on the
outer flank. The apertural branch forms a rather high ventro-
lateral projection, and forms a shallow sinus on the venter.
The other, smaller specimens are similar in conch shape and
ornament. Some of them, such as specimen MB.C.19128.2
(7 mm dm) and MB.C.19128.5 (4.2 mm dm) possess constric-
tions extending parallel to the riblets (Table 1). The tetrangu-
larly coiled stage ends at about 6 mm conch diameter.
R e m a r k s : The species is the type species of the subge-
nus Gastrioceras (Branneroceratoides) Kullmann (in Ste-
vanović & Kullmann 1962), which therefore has to be put in
synonymy with Entogonites.
E. tetragonus differs from E. grimmeri in the poorer devel-
opment of the tetrangular coiling of the inner whorls. The rib-
Fig. 5. Entogonites tetragonus (Kullmann, 1962) from sample DRZ 1
at Milivojevića Kamenjar section in Družetić; all ×5. A – Speci-
men MB.C.19128.1. B – Specimen MB.C.19128.2. C – Specimen
MB.C.19128.3. D – Specimen MB.C.19128.4. E – Specimen
KORN, JOVANOVIĆ, NOVAK and SUDAR
lets in E. tetragonus are, in comparison to E. grimmeri, a lot
less sharp. E. nasutus has also only weakly tetrangular inner
whorls, but has a much narrower umbilicus than E. tetrago-
nus, E. borealis has a narrower umbilicate than E. tetragonus,
and E. saharensis possesses sharper ribs with higher ventrolat-
Genus: Ubites gen. nov.
D e r i v a t i o n o f n a m e: After the River Ub in NW Serbia.
T y p e s p e c i e s : Ubites filipovici gen. nov. et sp. nov.
G e n u s d e f i n i t i o n : Entogonitidae with complex ontog-
eny. Inner whorls evolute, adult whorls with narrower umbi-
licus caused by stronger whorl overlap. Inner whorls
filipovici: Ubites filipovici gen. nov. et sp. nov.; NW Serbia
pseudocyclus: Nomismoceras pseudocyclus Campbell,
Brown & Coleman, 1983, p. 97; Queensland.
Table 1: Conch dimensions (in mm) and proportions for reference specimens of Entogonites tetragonus (Kullmann, 1962).
dm ww wh uw ah
3.88 2.22 0.36 1.23 0.46 1.83 0.10
3.41 – 0.40 1.41 0.49 –
3.31 1.77 0.37 1.28 0.50 1.86 0.08
2.69 1.35 0.45 1.59 0.46 1.69 0.18
2.44 – 0.42 1.35 0.43 –
2.35 0.99 0.42 1.50 0.49 1.58 0.27
1.97 0.92 0.40 1.36 0.47 1.65 0.25
0.98 0.53 0.44 1.38 0.43 1.71 0.26
R e m a r k s : Ubites gen. nov. differs from Entogonites in
the normally coiled inner whorls and in the ontogenetic de-
velopment, which shows an adult stage in which the uw/dm
ratio is significantly reduced.
Ubites filipovici gen. nov. et sp. nov.
1962 Gastrioceras (Branneroceras) branneri branneri Smith – Kull-
mann in Stevanović & Kullmann, p. 86—87, pl. 2, fig. 1
D e r i v a t i o n o f n a m e : After Dr. Ivan Filipović, in rec-
ognition of his contribution to the geology and stratigraphy
of the Paleozoic rocks in Serbia.
H o l o t y p e : Specimen MB.C.19130.1 (Museum für
Natürkunde, Berlin), illustrated in Fig. 6A.
T y p e l o c a l i t y a n d h o r i z o n : Milivojevića Kamenjar
section, Družetić (NW Serbia); sample DRZ 2 (“upper hori-
zon”), Entogonites Genus Zone.
M a t e r i a l : 35 specimens; conch diameter up to 16.5 mm.
Fig. 6. Ubites filipovici gen. nov. et sp. nov. from sample DRZ 2 (A—E) and sample DRZ 1 (F) at Milivojevića Kamenjar section in Družetić;
all 5. A – Holotype MB.C.19130.1. B – Paratype MB.C.19130.2. C – Paratype MB.C.19130.3. D – Paratype MB.C.19130.4. E – Paratype
MB.C.19130.5. F – Paratype MB.C.19129.
EARLY LATE VISEAN AMMONOID FAUNAS (NW SERBIA)
D i a g n o s i s : Species of Ubites gen. nov. with a thinly dis-
coidal, evolute conch (ww/dm = 0.35—0.45; uw/dm = 0.50—0.55)
at 8 mm dm; conch thinly discoidal, evolute (ww/dm =0.40;
uw/dm =0.47) at 16 mm dm. Aperture low (WER = 1.50—1.60)
in the adult stage. Shell ornament sharp biconvex riblets; inner
whorls with sharp dichotomizing ribs. Ventrolateral projec-
tion of riblets low in the adult stage, ventral sinus moder-
D e s c r i p t i o n : Holotype MB.C.19130.1 is an incom-
plete, slightly distorted specimen with 16 mm conch diame-
ter (Table 2); it demonstrates a rapid ontogenetic change
from the evolute juvenile stage into the adult morphology.
This change takes place within only one volution, in which
the whorl height becomes significantly larger. At 16 mm dm,
the flanks are slightly flattened and the umbilical margin is
subangular. The last volution shows the excellently pre-
served shell ornament, consisting of sharp riblets, of which
three stand within one millimeter. Every second of these rib-
lets is intercalated near the umbilical margin; the main riblets
begin on the umbilical wall and form a sharp elongate node
on the umbilical margin. The riblets form a shallow sinus on
the umbilical margin, a low projection in the inner flank
area, a very shallow sinus on the midflank, a low ventrolater-
al projection, and a moderately deep ventral sinus. Parallel to
the growth lines extend two shell constrictions standing ap-
proximately 120° apart. They begin in the inner flank area
and extend with the same depth across flanks and venter.
The smaller paratype MB.C.19130.2 (8.1 mm conch diam-
eter) represents the juvenile growth stage, in which the
conch is still evolute (uw/dm = 0.51). The whorls are wide
(ww/wh = 1.42) and crescent-shaped in this stage, and flanks
and venter are regularly rounded. The shell ornament shows
about 50 sharp riblets; between these, finer intercalatory rib-
lets begin in the inner flank area. The riblets extend almost
linearly across the inner flanks, bend forward to form a mod-
erately high ventrolateral projection, and then turn back for a
moderately deep ventral sinus.
Small specimens such as paratype MB.C.19130.4 (5.6 mm
conch diameter) display a similar conch morphology and
shell ornament. This specimen is more widely umbilicate
(uw/dm =0.57) than the larger ones.
R e m a r k s : Kullmann (in Stevanović & Kullmann 1962)
had only two small specimens, and he attributed these, be-
cause of the dichotomizing ribs, to the North American Mor-
rowan (late Bashkirian) index species Branneroceras
branneri. This determination did not consider the small size
of the Serbian specimens, which superficially resemble juve-
niles of Branneroceras branneri. The reconstruction of the
suture line by J. Kullmann (op. cit., fig. 4a) cannot be con-
firmed, as the specimen does not show the suture line.
The new species is similar to U. pseudocyclus (Campbell,
Brown & Coleman, 1983), but that species has, at 16 mm di-
ameter (i.e. the same size as the holotype of U. filipovici gen.
nov. et sp. nov.), less sharp riblets with a more linear course
across flanks and venter. The umbilicus is wider in U.
pseudocyclus (uw/dm = 0.47, in contrast to 0.42 in U. filipov-
ici gen. nov. et sp. nov.). U. pseudocyclus co-occurs, at the
type locality Mundubbera (Queensland), with species of the
genera Irinoceras, Bollandites, Maxigoniatites, and Cant-
abricanites, which can be taken as evidence for an early late
Visean age such as U. filipovici gen. nov. et sp. nov.
Acknowledgments: We greatly acknowledge the support for
the research program. D. Korn was supported by the Deut-
sche Forschungsgemeinschaft (DFG Project KO 1829/8-1).
The research of D. Jovanović and M. Sudar was supported
by the Ministry of Science and Technological Development
of the Republic of Serbia, Project No. 146009. We would
particularly like to thank Ivan Filipović for his advice in the
field. The manuscript profited from the reading by Sonny
Walton (Potsdam). We are indepted to J. Hladil, I. Filipović
and T. Lehotský for rewieving the typescript.
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dm ww wh uw ah ww/dm
16.35 6.91 5.12 7.12 3.41 0.42 1.35 0.44 1.60 0.33
8.91 3.35 2.29 4.64 1.63 0.38 1.46 0.52 1.50 0.29
8.08 3.28 2.31 4.15 1.58 0.41 1.42 0.51 1.55 0.32
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– 0.55 1.47 0.30
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2.56 1.41 3.01 0.94 0.47 1.82 0.55 1.46 0.33
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