2016 | 69/2 | 195–200 | 2 Figs. | 1 Pl. | www.geologia-croatica
 Journal of the Croatian Geological Survey 
and the Croatian Geological Society 

1. INTRODUCTION
The Scaphopoda is a class of Mollusca, also known as the “tusk 
shells”. Their shell is tubular and generally curved, having open­
ings at both ends. Scaphopods are a cosmopolitan marine infau­
nal species, and live from the intertidal zones down to depths of 
more than 6000 m. Scaphopod species mostly burrow in sand or 
mud, with their head pointed downwards, and the narrow poste­
rior end of the shell protruding above the substratum (KNUD­
SEN, 1964; LAMPRELL & HEALY, 1998 and references therein; 
HABDIJA et al., 2011).

Although the scaphopod systematics have been revised 
over a few decades, the time of their appearance is still a matter 
of debate, as are their phylogenetics, so they are considered to be 
a puzzling molluscan class. Two orders of scaphopods are 
known: Dentaliida and Gadilida, which differ both morphologi­
cally, in shell and foot form, and behaviourally, in different bur­
rowing activities (LAMPRELL & HEALY, 1998 and references 
therein; STEINER & KABAT, 2001, 2004).

This paper focuses on the Badenian dentaliid species Fissi­
dentalium badense (PARTSCH in HÖRNES), recorded in the 
Mio cene deposits of the Paratethys, and widely distributed in Ba­
denian sediments of the Central Paratethys (e.g. BALDI, 1960; 
KOJUMDGIEVA & STRACHIMIROV, 1960; BALLUK, 1972; 
TITA, 2007; HARZHAUSER et al., 2011). Specimens of F. ba­
dense included here were collected from the Badenian sediments 
of the south and southwestern margin of the Central Paratethys 
(Croatia, Bosnia and Herzegovina and Serbia). The analyzed spe­
cimens are housed at the Natural History Museum Belgrade 
(NHMB) and Croatian Natural History Museum (CNHM). The 
first finding of F. badense from Bosnia and Herzegovina is pub­
lished here.

2. GEOLOGICAL SETTING
The Central Paratethys extended from Bavaria to the Carpathian 
mountain chain (e.g. RÖGL 1998, 1999). Badenian sediments 

 
Fissidentalium badense (PARTSCH in HÖRNES, 1856) from 
the Badenian deposits of the south and southwestern margin  
of the Pannonian Basin System (Central Paratethys)
Gordana Jovanović1 and Marija Bošnjak2

1 Natural History Museum Belgrade, Department of Paleontology, Division of Geology, Njegoševa 51, Belgrade, Serbia; (gordana.j@nhmbeo.rs)
2 Croatian Natural History Museum, Department of Geology and Paleontology, Demetrova 1, Zagreb, Croatia; (corresponding author: marija.bosnjak@hpm.hr)

doi: 10.4154/gc.2016.20     

Abstract
The Badenian (Middle Miocene) Scaphopoda from the south and southwestern margin of the 
Pannonian Basin System (Central Paratethys) are poorly documented, and in published papers 
mostly presented by short lists of identified species. For the present study, 300 scaphopod speci-
mens were examined with the focus on a singular species, Fissidentalium badense (PARTSCH 
in HÖRNES, 1856). This species is represented by 152 specimens housed in the Natural His-
tory Museum Belgrade and the Croatian Natural History Museum in Zagreb. The most numero-
us and the best preserved specimens of F. badense were collected at the Višnjica locality near 
Belgrade (Serbia), while the others originate from the vicinity of Zagreb (Croatia) and Ugljevik 
(Bosnia and Herzegovina).

from the investigated area belong palaeogeographically to the 
south and southwestern margin of the Central Paratethys, and 
geotectonically to the Pannonian Basin System, surrounded by 
the Alps, Carpathians and Dinarides (Fig. 1) (e.g. RÖGL 1998, 
1999; PAVELIĆ, 2001, 2002; PILLER et al., 2007).

The beginning of the Badenian epoch is marked by a marine 
transgression, which flooded the areas behind the newly uplifted 
mountain chains (Paratethys) and was also prominent in the Medi­
terranean (Fig. 1) (RÖGL, 1998, 1999; KOVÁČ et al., 2007). Ac­
cording to numerous papers, the Badenian deposits of the south 
and southwestern margin of the Central Paratethys lie discorda­
ntly on the older Miocene, Mesozoic or even Palaeozoic deposits. 
The contact with basal rocks is in many places unclear or erosive 
or tectonic in character (e.g. ŠIKIĆ et al., 1978, 1979; BASCH, 
1983a, b; VRABAC & MIHAJLOVIĆ, 1990; ANĐEL KOVIĆ et 
al., 1991; PAVELIĆ, 2001, 2002; SAVIĆ et al., 2005; PIKIJA, 
2009).

At the southern margin of Central Paratethys, F. badense is 
recorded in the Lower Badenian deposits of Serbia (wider Bel­
grade region) (PETKOVIĆ et al., 1976), in the assumed Lower to 
Middle Badenian of western Serbia (ŽUJOVIĆ, 1889; PAVLOVIĆ, 
1903; STEVANOVIĆ, 1949; PETROVIĆ, 1967), and in the Lower 
Badenian (Upper Lagenidae zone) in northeastern Bosnia and 
Herzegovina (Bogutovo Selo) (VRABAC et al., 2013, 2014; PE­
ZELJ et al., 2013) (Fig. 1). In the area of the southwestern margin 
of the Central Paratethys, this species is present in the Miocene 
(probably Lower Badenian) marls of the wider Zagreb region 
(Med vednica Mt., Čučerje area) (KOCHANSKY, 1944, 1957; 
ĆORIĆ et al., 2009), and in the assumed Middle to Upper Bade­
nian deposits of the Samobor area (Vrhovčak locality, previously 
known as „Zaprešić Breg“ or „Zaprešić brijeg“) (ŠUKLJE, 1929; 
PAVLOVSKY, 1957, 1960; BAJRAKTAREVIĆ, 1978; VRSALJ KO, 
2003; BAKRAČ et al., 2010). In the Badenian sediments of the 
Glinsko Pokuplje area, this species is found at one locality (PI­
LAR, 1873).

Article history:

Manuscript received December 16, 2015 
Revised manuscript accepted May 31, 2016 
Available online June 28, 2016 
 
 

Keywords: Fissidentalium badense, Badenian, 
Pannonian Basin System, Central Paratethys



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3. MATERIALS AND METHODS
Three hundred scaphopod shells were examined, and 152 of them 
belong to the species Fissidentalium badense (PARTSCH in 
HÖRNES). All the analyzed specimens are damaged, mostly in 
the apex area, and the majority of specimens are preserved as 
shell fragments.

NHMB specimens were collected at the Višnjica locality 
(wider area of Belgrade), and they are a part of the P. Pavlović 
collection, assembled in 1897 (90 specimens) and the P. Ste va­
nović collection, assembled in 1962 (40 specimens). Other spec­
imens come from the Loznica area in western Serbia (3 speci­
mens), and Bogutovo Selo near Ugljevik, Bosnia and Herzegovina 
(1 specimen).

CNHM specimens originating from the wider Zagreb region 
are part of the „Marine Miocene fauna of the southern part of the 
Medvednica Mt.“ collection (10 specimens) and the „Miocene 
fauna of Zaprešić Breg near Samobor“ collection (5 specimens). 
Specimens from the third locality are part of the „Miocene and 
Pliocene fauna from the Glinsko Pokuplje area and Zrinjsko­
Dvorska valley“ collection (3 specimens).

The site yielding the largest number of scaphopods is 
Višnjica (Fig. 1), with 280 specimens from clayey sediments, 130 
among them belonging to F. badense (LUKOVIĆ, 1922; JOVA­
NOVIĆ & JOVANOVIĆ, 1998). This fossil fauna is the best pre­
served, and therefore chosen to be described here.

Determination of species is based upon several papers (e.g. 
HÖRNES, 1856; SACCO, 1897; KOJUMDGIEVA & STRA­
CHIMIROV, 1960; PAVIA, 1991; STEINER & KABAT 2001, 
2004). Parameters used in the determination are after STEINER 
(1999) and SILVA­FILHO et al. (2012) (Fig. 2).

4. RESULTS
SYSTEMATIC PART
Class Scaphopoda Bronn, 1862
Order Dentaliida da Costa, 1776
Family Dentaliidae Children, 1834
Genus Fissidentalium Fischer, 1885
Subgenus Antalis Adams & Adams, 1854

Fissidentalium badense (Partsch in Hörnes, 1856)
(Pl. 1, Figs. 1­6)

Type species: Dentalium ergasticim FISCHER, 1885, monotype
1856 Dentalium Badense – HÖRNES, p. 652, pl. 50, fig. 30
1897 Entalis badensis (PARTSCH) et var. – SACCO, p. 107–108, 

pl. 9, figs. 17–30
1925 Dentalium (Entalis) badense PARTSCH var. borealis 

KAUTSKY – KAUTSKY, p. 53–54, pl. 5, fig. 12
1929 Dentalium badense (PARTSCH) – ŠUKLJE, p. 52
1947 Dentalium (Fissidentalium) badense PARTSCH – TAU­

BER, p. 302, pl. 7, figs. 8–10, pl. 8, fig. 3
1897 Entalis badensis – SACCO, p.107, pl. 9, figs. 17–20
1960 Dentalium (Entalis) badensis PARTSCH in HÖRNES – 

KOJUMDGIEVA & STRACHIMIROV, p. 225, pl. 52, figs. 
19–20

1960 Dentalium badense PARTSCH – BALDI, p. 57, pl. 1, fig. 5
1991 Fissidentalium badense (PARTSCH in HÖRNES, 1856) – 

PAVIA, p. 146, pl. 5, fig. 4; pl. 6, fig. 6. (cum syn.).
1998 Dentalium badense PARTSCHI – JOVANOVIĆ & 

JOVANOVIĆ, p. 201–202
2011 Fissidentalium badense (PARTSCH in HÖRNES, 1856) – 

HARZHAUSER et al., p. 219, pl. 8. figs. 6–7
2013 Dentalium (Antalis) badense (PARTSCH, 1856) – ROUT­

NEROVÁ, p. 20, pl. 2, fig. 2

Material: 20 relatively well preserved and 110 damaged spe­
cimens from the Višnjica locality, 3 specimens from the Loznica 
area (Serbia), 10 damaged specimens from the Medvednica Mt. 
(Čučerje area), 5 specimens from the „Zaprešić Breg“ locality and 
3 specimens from the Glinsko Pokuplje area (Croatia), 1 specimen 
from Bogutovo Selo near Ugljevik (Bosnia and Herzego vina).

Dimensions: L = 37.3 mm; Larc = 32.0 mm; Ha = 6.4 mm; 
Wa = 6.3 mm; Hp = 2.9 mm; Wp = 2.8 mm.

Description: Shell is medium to large, solid, moderately 
curv ed, narrow and elongated, open at both ends. Dorsal side is 
concave, and the ventral is convex. The apex is damaged in all 
the collected specimens. The shell narrows and slightly curves to 
the wider anterior opening. The shell is ornamented with solid 
radial ribs and growth lines. Sharp and moderately high, 8 to 10 
radial ribs are near the apical area. Ribs become lower and almost 
flat near the wider end of the shell. Secondary ribs are inserted 

Figure 1. Geographic and paleogeographic setting of the investigated area  with 
marked localities containing recorded dentaliids (after KOVÁČ et al., 2007).

Figure 2. Morphometric measurement parameters: L − length, Max – maximum 
diameter, Dmax – maximum distance from the anterior opening, Arc – curvatu-
re maximum, Larc − maximum length from the apex to the deepest point of the 
curvature, Ha − height of the ventral opening, Wa − width of the ventral ope-
ning, Hp − apical height, Wp − apical width (after STEINER, 1999; SILVA-FILHO 
et al., 2012).



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Gordana Jovanović and Marija Bošnjak: Fissidentalium badense (PARTSCH in HÖRNES, 1856) from the Badenian deposits of ... 197

Plate 1. Fissidentalium badense (PARTSCH in HÖRNES, 1856).

1 Višnjica locality, Serbia (Inv. No. K−2347/1);
2 Loznica, Serbia (Inv. No. K−2689);
3 Bogutovo Selo, Bosnia and Herzegovina (Inv. No. K−6746);
4 Čučerje, Croatia (Inv. No. 573); 
5 Vrhovčak („Zaprešić Breg“), Croatia (Inv. No. 1.413); 
6 Glinsko Pokuplje area, Croatia (Inv. No. 1.220).



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between the primary ribs at various distances from the apex. The 
maximum number of inserted ribs near the posterior part of the 
shell does not exceed 30. The primary and secondary ribs at the 
posterior side of the shell are almost indistinguishable from each 
other with only minor differences. The species varies in number 
and thickness of the secondary ribs. Concave interrib spaces vary 
in width. Together with the radial ribs, fine and dense irregular 
growth lines (except in the apical area) characterize the shell sur­
face. The cross­section of both ends is subcircular.

Remarks: Many authors studied the phylogeny and classifica­
tion of the scaphopods (e.g. REYNOLDS, 1997; REYNOLDS & 
AKOKO, 1999). Fissidentalium badense (PARTSCH in HÖRNES, 
1856) was first recognized as Dentalium badense (HÖRNES, 1856), 
Entalis (SACCO, 1897) or Dentalium (Entalis) badensis (KOJUM­
DGIEVA & STRACHIMIROV, 1960). F. badense from the Višnjica 
locality, and presented here, fits the descriptions by PAVIA (1991). 
The species described in KOJUMDGIEVA & STRACHIMIROV 
(1960) has more secondary ribs (35), as do several specimens from 
Koritnica, Poland (BALLUK, 1972). STEINER & KABAT (2004) 
single out the subgenus Dentalium LINNAEUS 1758. In the re­
search of the taxonomy and ecology of the recent genus Fissiden­
talium (FISCHER), LAMPRELL & HEALY (1998) indicate the 
significance of the aperture, and the number and rib morphology 
that can contribute to the research of the fossil species.

Geographic and stratigraphic distribution: Fissidenta­
lium badense (PARTSCH in HÖRNES, 1856) appears during the 
Burdigalian in the Mediterranean at the Torino Hills (SACCO, 
1897), and in the Lower Miocene of the Paratethys (HARZHAUSER, 
2002; HARZHAUSER et al., 2011). F. badense has a wide geogra­
phic distribution, and represents a species typical of the Middle 
Miocene of the Paratethys (e.g. KOJUMDGIEVA & STRACHI­
MIROV, 1960; PAVIA, 1991; HARZHAUSER et al., 2011).

5. DISCUSSION
In the Badenian sediments of the south and southwestern margin 
of the Central Paratethys, the majority of scaphopod discoveries 
belong to the dentaliids. Here, we analyzed and revised Fissiden­
talium badense (PARTSCH in HÖRNES, 1856) specimens stored 
at the NHMB and CNHM. Given the number of the recorded 
dentaliids, Višnjica near Belgrade (Serbia) is the site yielding the 
largest number of specimens.

At the Višnjica locality, scaphopods were discovered in the 
clayey sediments, known as the „Višnjica clays“ or the „clays with 
Pleurothoma“. This site was compared to the Badenian stratotype 
Baden­Sooss in Vienna (PAVLOVIĆ, 1903; LUKOVIĆ, 1922), 
which contains many specimens of F. badense. The stratigraphic 
position of the „Višnjica clays“ is not precisely determined. Based 
upon foraminifera, STEVANOVIĆ (1977) attributes it to the Mid­
dle Badenian, and GRUJIČIĆ (2010) specifies the zone with Spi­
ro rutilus carinatus (d’ORBIGNY). Sandstones containing rare 
fine­shelled pectens (Costelamussium sp.), Aequipecten macrotis 
(SOWERBY) and Aequipecten scabrella (LAMARCK) (LUKO­
VIĆ, 1922) cover the clay stratum. These pectinid species also 
occur in the Lower Badenian of Bukovac on the Fruška gora Mt. 
(JOVANOVIĆ, 2014), in the Grund Gaindorf formation in Aus­
tria (MANDIC & HARZHAUSER, 2003; MANDIC, 2004), and 
in the Lower Badenian (Lagenidae zone) sediments of Szokoly, 
Hungary (BALDI, 1960). The „Višnjica clays“ contain numerous 
scaphopod specimens: Fissidentalium badense (PARTSCH in 
HÖRNES), Gadilina jani HÖRNES, Dentalium sexangulum 
GMELIN, Dentalium (Antale) novemcostatum decemcostulata 

SACCO, Dentalium (Dentalium) michelottii HÖRNES and Fis­
sidentalium mutabile (DODERLEIN).

In the Lower Badenian argillaceous marls of the wider Bel­
grade region, deep sea molluscs are very common: Neopycnodonte 
cochlear POLI, Lentipecten corneus denudatus (REUSS), Vaginella 
austriaca KITTL and Aturia aturi (BASTEROT) (PET KOVIĆ et 
al., 1976; JOVANOVIĆ, 2014). These findings, as well as the record 
of Parvamussium duodecimlamellatum (BRONN), indicate a 
Lower Badenian age (STUDENCKA et al., 1998; SELMECZI et 
al., 2012; JOVANOVIĆ, 2014). In the Miocene (probably Lower 
Badenian) marls of the wider Zagreb region („Čučerje“ develop­
ment on the Medvednica Mt.), the wider Belgrade region and Ug­
ljevik (Bogutovo Selo), together with dentaliids, scaphopods are 
accompanied by a pelagic fauna of nautilids (Aturia aturi (BAS­
TEROT), pteropods (Vaginella austriaca KITTL) and planktonic 
foraminifers (KOCHANSKY, 1944; STEVANOVIĆ, 1977, AVA­
NIĆ et al., 1995;  JOVANOVIĆ, 2014). These known data, together 
with further detailed dentaliid analysis, can contribute to the re­
construction of the first Middle Miocene marine transgression in 
the south and southwestern margin of the Central Paratethys.

All the published data and records suggest that F. badense 
mostly lived in muddy and muddy­sandy substrate at all the stud­
ied localities in Serbia, Bosnia and Herzegovina and Croatia. This 
species was widespread during the Lower Badenian, with fossil 
evidence from the clayey sediments of Austria (Baden­Sooss) 
(RÖGL et al., 2009), Romania (Lăpugiu de Sus) (TITA, 2007), 
Slovakia (BALLUK, 1972; HARZHAUSER et al., 2011), Hungary 
(BALDI, 1960) and Bulgaria (KOJUMDGIEVA & STRACHIMI­
ROV, 1960). The distribution of F. badense indicates the contem­
porary marine connections and migration routes. Due to the re­
corded and afore­mentioned findings, possible migration routes 
of F. badense to the area of Paratethys could be the southwestern 
(„Transtethyan corridor“) and the northern marine passages (e.g. 
RÖGL, 1998; STUDENCKA et al., 1998; KOVÁČ et al., 2007).

6. CONCLUSIONS
This paper presents the scaphopod species Fissidentalium ba dense 
(PARTSCH in HÖRNES, 1856) housed at the Natural History 
Museum Belgrade and the Croatian Natural History Museum. 
Fossil material was collected from the Badenian (Middle Mio­
cene) deposits of the south and southwestern margin of the Cen­
tral Paratethys.

The age of the analyzed specimens from Serbia and Bosnia 
and Herzegovina stored at the Natural History Museum Belgrade 
is the Lower Badenian, while other Middle Miocene findings are 
questionable in age and require further research.

During the upper part of the Lower Badenian, the species 
had a wide geographic distribution, which implys an open marine 
connection of the Central Paratethys with other marine areas.

According to the available fossil record, Fissidentalium 
badense lived on a muddy to muddy­sandy substrate.

ACKNOWLEDGMENT
The authors thank Prof. Sejfudin VRABAC (Faculty of Mining, 
Geology and Civil Engineering, University of Tuzla, Bosnia and 
Herzegovina), Prof. Jasenka SREMAC (Faculty of Science, Uni­
versity of Zagreb, Croatia), Dr. Davor VRSALJKO (Croatian Na­
tural History Museum, Zagreb) and anonymous reviewers for 
useful comments which helped us to improve the paper, and to 
Dr. Davorka RADOVČIĆ (Croatian Natural History Museum, 
Zagreb) for help in translating the manuscript.



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