Middle Miocene Shark Teeth from the Southern Margin of the Pannonian Basin System (Serbia, Central Paratethys)

GEOLOŠKI ANALI BALKANSKOGA POLUOSTRVA Volume 80 (1), July 2019, 29–43 – DOI: 10.2298/GABP1901029J

Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

GORDANA JOVANOVIĆ1, N ICOLAE TRIF2, VLAD CODREA3 & D RAGANA ĐURIĆ1

Abstract. This paper describes Middle Miocene (Badenian) shark teeth from Serbia and discusses their geographical distribution at other localities of the Central Paratethys. The shark teeth originate from Višnjica (near Belgrade), from Višnjica Clay, or the ‘Pleurotoma Clay’. The variety of sharks is very low, but according to these fossils Višnjica is the richest among Serbian localities. The sharks teeth reported in the paper belong to the following taxa: Otodus (Megaselachus) megalodon (A GASSIZ, 1835), Hemipristis serra AGASSIZ, 1835 and Odontaspididae indet. The high diversity of invertebrates (molluscs, echinoids, Key words: corals etc.) and other coeval fossil assemblages indicate a warm period (the Middle Miocene, Badenian, Middle Miocene Climatic Optimum), which preceded the Middle Miocene Cli- sharks, Paratethys Sea, Serbia. matic Transition.

Апстракт. У овом чланку сe описују средњомиоценски (баденски) зуби ајкула из Србије и дискутује о њиховој географској дистрибуцији на другим локалитетима Централног Паратетиса. Зуби ајкула потичу из Вишњице (недалеко од Београда), из Вишњичких глина или “Плеуротом- ских глина”. Разноврсност ајкула је врло ниска, али је по овим фосилима Вишњица најбогатија међу српским локалитетима. Описани зуби ајкула припадају следећим таксономима: Otodus (Megaselachus) megalodon (A GASSIZ, 1835), Hemipristis serra AGASSIZ, 1835, и Odontaspididae indet. Кључне речи: Висока разноврсност бескичмењака (мекушаца, ехиноида, корала) као и Средњи миоцен, баден, других фосилних заједница указују на топли период (средњомиоценски ajкуле, море Паратетис, климатски оптимум), који је претходио средњомиоценској климатској Србија. транзицији.

1 Natural History Museum, Njegoševa 51, 11000 Belgrade, Serbia. E-mails: [email protected];[email protected] 2 Brukenthal National Museum, Natural History Museum, Sibiu, Romania, Cetăţii Street 1, Sibiu, RO-550160, Romania. E-mail: [email protected] 3 Babeș-Bolyai University, Faculty of Biology-Geology, Department of Geology 1, Kogălniceanu Str., Cluj-Napoca, RO-400084, Romania. E-mail: [email protected]

29 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

Introduction (1958), ANĐELKOVIĆ et al. (1989) and JOVANOVIĆ & TOMIĆ (1997). In the past decades, only a few new data have Miocene shark teeth from Paratethys are men- been added to this topic. tioned in the literature since the 19th century with This paper refers to the Badenian shark fauna numerous described extinct species (e.g., AGASSIZ, from Višnjica (Belgrade, Serbia) (Fig. 1A, B, C). The 1843; PROBST, 1878; KOCH, 1903; SCHULTZ, 1971, 1979, aim of this study is to increase the knowledge about 2013). Recent research allows the evaluation of their the shark diversity and its distribution along the diversity, as well as the areal and the stratigraphic southern margin of the Central Paratethys during the distribution of these fossils. Many references deal Middle Miocene. This study of the shark teeth pills a with the taxonomy and the reassessment of these gap within this region, allowing correlations be- sharks (e.g. CAPETTA, 2012; MIKUŽ & ŠOSTER, 2013; tween Serbian fauna and the coeval well-docu- MIKUŽ et al., 2014, 2015, 2017; SZABÓ & KOCSIS, 2016; mented similar faunas reported from the various TRIF et al., 2016; TRIF & CODREA, 2017). The present other areas of the Paratethys. By analysis of the Mid- analysis points out that the shark teeth are relatively dle Miocene Serbian localities, the Višnjica fauna ap- rare in the palaeontological record of the Serbian pears to be the most diverse. In addition to the taxa Miocene localities; but it also replects a broad geo- previously reported herein, GRUBIĆ (1958) mentioned graphic distribution in the Central Paratethys Sea. the following species: Carcharodon megalodon AGAS- SIZ, Lamna cuspidata AGASSIZ and Lamna (Otodus) appen diculata AGASSIZ. The last species is one of the most frequently reported nominal shark species from the Creta- ceous and Paleocene according to SIVERSSON et al. (2015) and CAPPETTA (2012), but not from the Miocene. Therefore, it is likely to be a misidenti- pied taxon. The Miocene shark teeth from other Serbian localities, like Rakovica and Go- lubac, have generally been assigned to Lamna, or just to ‘pish teeth’ (MIKINČIĆ, 1932; Fig. 1. Location of Višnjica locality in: A. Europe; B. Serbia; C. Belgrade area. JOVANOVIĆ & TOMIĆ, 1997). With the exception of a single specimen of Odontaspididae from The fossil shark teeth from Serbia have been Rakovica, no other shark teeth could be found in the poorly studied in comparison with other fossils (e.g. collections of the Belgrade Museum. ĐURIĆ et al., 2017: RADOVIĆ & BRADIĆ-MILINKOVIĆ, 2018 ). The Middle Miocene (Badenian) sediments from Serbia are exposed in various regions of the country. Geological setting These sediments are rich in fossils, mainly inverte- brate fossils. Shark teeth are mentioned only in a In the Paleogene, the collision between the small number of localities, such as Višnjica and Rako- African and the European plates has resulted in divi- vica, from the vicinity of Belgrade, and in eastern Ser- sion of the Tethyan Realm into two different paleo- bia (Golubac), according to MIKINČIĆ (1932), GRUBIĆ geographical realms - the circum-Mediterranean in

30 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

the south and the Paratethys in the north (LASKAREV, The Badenian age of the deposits near Belgrade 1924; RÖGL & STEININGER, 1984; RÖGL, 1998; POPOV et is based on the study of foraminiferal assemblages. al., 2004). On its turn, the Paratethys can be divided It comprises three units: the Early Badenian (the La- into three distinct regions: Western, Central, and genidae Zone), the Middle Badenian (the Spirorutilus Eastern Paratethys (PILLER et al., 2007). Each region carinatus Zone) and the Late Badenian (Elphidium had specipic developments in terms of facies and dis- crispum and Ammonia beccarii Zone) (PETROVIĆ, 1970;

Fig. 2. A. Geological map of the Višnjica area (after the map 1:100.000 of Serbia, sheet Pančevo, according to IVKOVIĆ et al. 1966, modiFied); B. The map of Central Paratethys (after RÖGL & S TEININGER, 1984, modiFied).

tribution. The former Central Paratethys region is 1985). The VCF exposures have been reported in the congruent with the area of nowadays Austria, Slove- Ramadan stream, at its conpluence with the Danube. nia, Poland, Hungary, Croatia, Serbia, Bosnia and The outcrops from the banks of the Danube studied Herzegovina, Ukraine and Romania. Several Middle in the 19th and 20th centuries allowed the collection Miocene sedimentary basins from Serbia have been of these fossils. However, after the construction of the closely connected to this area, which allowed faunal dam at hydroelectric power plant “Đerdap” (Iron exchanges. In the Middle Miocene, in the vicinity of Gates), the Danube level rose and these outcrops are Belgrade, the Paratethys Sea plooded different types currently plooded. of basement and a normal marine regime occurred in the area. The related sedimentary deposits are very rich in fossils. The pirst geological studies in the Material and methods area enabled a large number of typical marine taxa to be reported. In the Višnjica site, the largest group This study is based on the review of pifteen fossil of taxa is represented by Badenian gastropods from shark teeth collected during several past pield works. the so-called ‘Pleurotoma Clay’, or ‘Višnjica Clay’ The studied shark teeth originate from three distinct (LUKOVIĆ, 1922; STEVANOVIĆ, 1977; ANĐELKOVIĆ et al., collections, all obtained from the same Badenian de- 1989) assigned to the Višnjica Clay Formation (ab- posits. The collectors were: Petar Pavlović, at the end breviated VCF). of the 19th century, Petar Stevanović, in 1959, and Ve-

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 31 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

limir Milošević, around the mid-20th century. Mor- well preserved, pine and regular. The neck is frag- phological features have been used to provide de- mented, but it can be noticed in the central part, on tailed taxonomic data. We considered the shape and the lingual side of the tooth. The lingual side is mode- size of the main cusp and the serration, where avail- rately convex, while the labial one is plat. The shape able. The photos of the specimens have been taken of the crown suggests an antero-lateral position of with an OLYMPUS Z4001 camera and a PANASONIC this tooth. DMC-FZ50 camera, respectively, attached to a BIOOP- The tooth in the Figs. 3h-j is also of a rather mod- TICA 100 stereomicroscope. est size, with a height of 61 mm and a preserved The shark teeth found in the VCF range in size width of 48 mm. The general shape is similar to the from less than 8 mm to over 80 mm. These have been other two teeth already described. The cutting edge collected directly by hand, without sieving the sedi- is strongly worn-out and the serration is only pre- ment. The fossils are housed at the Natural History served in a small portion. It is interesting to note the Museum in Belgrade, Serbia (abbreviated as NHMB; bio-erosion marks on the enamel of this tooth. Both collection numbers: K 7050; PS 23-26; KV-P 282- sides of the crown bear hundreds of indentations 291). Some of the shark teeth are poorly preserved caused by an unknown organism. Based on the gene- and eroded with exposed enamel cracks. The roots ral shape, it is likely that this tooth had an upper an- are damaged too. This state of preservation suggests terior position. that some teeth have been exposed to wave action, Discussions. The taxonomy of the Otodontidae which led to fragmentations, abrasion and polishing. family has been debated for a long time, particularly Most of the teeth are black and dark grey, while oth- over the last decade (JORDAN & HANNIBAL, 1923; GLICK- ers are brownish or even white. MAN, 1964; SCHULTZ, 1971; PIMIENTO et al., 2010; CAP- PETTA, 2012 etc.). Based on the presence, absence or size of the serrations and cusplets, as well as on the Systematic paleontology differences in the root morphology, CAPPETTA (2012) revised and divided the Otodus genus into three sub- Superorder Galeomorphii COMPAGNO, 1973 genera. The described morphology allows us to con- Order Lamniformes BERG, 1958 pidently assign the specimens to the Otodus Family Otodontidae GLICKMAN, 1964 (Megaselachus) megalodon species (CAPPETTA, 2012). Genus Otodus AGASSIZ 1843 (sensu CAPPETTA, 2012). Although this species has a cosmopolitan distribution and it is by far better known from the Mio- Otodus (Megaselachus) megalodon AGASSIZ, 1835 Pliocene formations of the North, Central and South America (FOWLER & KUMMEL, 1911; DE MUIZON & Material: Three teeth (Figs. 3.a-d; e-g and h-j); DEVRIES, 1985; PIMIENTO et al., 2013; CARRILLO-BRICENO NHMB: PS 23, K 7050, PS 24. et al., 2015), it is also ubiquitous in the Central The tooth illustrated in Fig. 3.a-d is the biggest Paratethys Sea. Otodus (M.) megalodon has been and the most complete tooth ever found in this local- found in the Badenian deposits from over 50 out- ity. It has a height of 82 mm and a width of 57 mm. crops in: Austria (ZAPFE, 1954; FLÜGEL & KOLLMANN, Its general shape is triangular with a strong root and 1964; KOLLMANN, 1969; SCHULTZ, 1971; HIDEN, 1995; a well depined neck. The lingual side is convex (Fig. RÖGL et al., 2008; SCHULTZ, 2013), the Czech Republic 3a), while the labial one is plat (Fig. 3b). The cutting (S CHULTZ et al., 2010), Hungary (SZABÓ & KOCSIS, 2016), edge is well preserved and has a pine and regular ser- Slovenia (MIKUŽ, 2000; MIKUŽ & ŠOSTER, 2013; MIKUŽ et ration (Fig. 1c). No cusplets are present at the base al., 2014, 2015), Croatia (PAUNOVIĆ, 1987), Poland of the crown. Based on the general shape of the tooth, (P AWLOWSKA, 1960; RADWANSKI, 1965; SCHUFLTZ, 1977, it most likely had an anterior position. 1979), Slovakia (HOLEC & SABOL, 1996; HOLEC, 2001) The tooth in the Figs. 3e-g is smaller, measuring and Romania (TRIF et al., 2016; TRIF & CODREA, 2017). only 48 mm in height and 37 mm in width. Most of Sometimes, associated teeth can also be found (see its root and the apex are missing. The serration is MIKUZ & ŠOSTER, 2013). It is interesting to note that O.

32 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

(M) megalodon is part of the fauna listed in the Odontaspididae family. These sharks had a wider dis- Badenian type-section of Baden-Sooss in Lower Aus- tribution during the Cenozoic than nowadays. As a tria (RÖGL et al., 2008). family of relatively large sharks, the Odontaspididae are known since the Aptian (early Cretaceous), ac- Order Carchariniformes COMPAGNO, 1973 cording to CIONE et al. (2007). Family Hemigaleidae HASSE, 1879 Genus Hemipristis AGASSIZ, 1843. Discussion Hemipristis serra AGASSIZ, 1835 Although the shark teeth from the Višnjica locality Material: One tooth (Figs. 3k-n); NHMB: PS 25 have been previously reported (GRUBIĆ, 1958), their The illustrated tooth is an upper anterior. The value has not been precisely considered in the strati- crown is bent distally, plat, with a small basal fold on graphic context. The stratigraphy of the VCF is still the labial side and convex on the lingual side. The debated, but all the available data indicate a Middle cutting edges are typical for H. serra with larger, rarer Miocene (Badenian, calcareous nannoplankton NN5 serrations on the distal side, and smaller and more Zone) age (Fig. 3). The sediments with shark teeth in numerous serrations on the mesial side. The apex is Serbia were deposited during the Early Middle Mio- devoid of any kind of serration. The root is only cene (Badenian) transgression. The Paratethys Sea partly preserved, while the mesial lobe is missing. plooded different terrains in Serbia, terrains which Discussions: Hemipristis serra is a common Mio- had either previously completely emerged, or had cene species, especially in the Badenian, where it has borne lacustrine sediments. It was a continuous ex- been found in a large number of outcrops across the panding transition from freshwater lake environ- Central Paratethys (SCHULTZ, 1977; BRZOBOHATÝ & ments to marine ones, on the whole southern sector SCHULTZ, 1978; SCHULTZ, 2013; SZABÓ & KOCSIS, 2016; of the Pannonian basin (ĆORIĆ et al., 2009; KRSTIĆ et TRIF & CODREA, 2017) and around the world (MÜLLER, al., 2012). Therefore, the diversity of the marine 1999; PURDY et al., 2001; CAPPETTA, 2012). Basically, it fauna from Višnjica as well as other Badenian locali- occurs in the same areas mentioned above for O. (M) ties originated mainly from migration events from megalodon, and it is particularly common in the the Mediterranean basin through the Slovenian cor- Vienna and Styrian basins. ridor. Connections are recorded also between the Pannonian basin and the Transylvanian one, through Family Odontaspididae MÜLLER AND HENLE, 1839 Mureș passageway (CHIRA & MĂRUNTEANU, 1999). In the Belgrade region, several Badenian litholo- Odontaspididae indet. gical units have been established such as the Rako- vica Sand, Višnjica Clay, Tašmajdan, Kalemegdan and Material: Seven teeth (one illustrated in Figs. Višnjica biogenic limestones (Leitha-type Lime- 3o-r); NHMB: KV-P 282 stone). Primary analyses were based on the forami- A small number of other shark teeth have been niferal and molluscan fauna. The foraminiferal fauna found in addition to the above described specimens. in Višnjica was studied by PAVLOVIĆ (1897) who They are all poorly preserved. The teeth are heavily pointed out its strong similarity to the one of the worn, most probably due to continuous wave action. ‘Baden Tegel’ (Vienna basin, Austria). Based on this The cutting edges and the apex are rounded and the rich fossil record, the Middle Miocene (Badenian) age lateral cusplets are reduced to just a few small was conpirmed in the vicinity of Belgrade (PAVLOVIĆ, bumps; the root is mostly fragmented, or totally 1922; LUKOVIĆ, 1922; GRUBIĆ, 1958). Later, foramini- missing, and in some cases polished to a smooth sur- fera and calcareous nannofossils were studied by face. However, the sigmoid propile and the slender PETROVIĆ (1970), STEVANOVIĆ (1977), MIHAJLOVIĆ & crowns that are characteristic for a tearing type of KNEŽEVIĆ (1989). GRUJIČIĆ (2010) pointed out the ex- dentition indicate that these teeth belong to the istence of the zone with Spirorutilus carinatus (D’OR-

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 33 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

fossil assemblages, the ﬁrst marine transgression in the Papuk Mt. (northern Croatia) according to ĆORIĆ et al., (2009) corresponds to the NN5 Zone. Recently, JOVANOVIĆ & BOŠNJAK (2016) based on the presence of some clams and scaphopods, assumed a late Early Badenian age. A main problem is related to the different interpretations of the Badenian stage subdivisions, as well as to the timing of Badenian marine transgressions in Paratethys (PILLER et al., 2007; KOVAČ et al., 2007; PEZELJ et al., 2013; HOHENEGGER et al., 2014; SANT et al., 2017). The different taphonomic conditions of the teeth suggest the origin from different stratigraphic levels. Additionally, a biostratigraphic revision is re- quired for some Badenian localities (JOVANOVIĆ et al., 2018). The sediments with shark teeth in Serbia were deposited during the early Middle Mio- cene (Badenian) transgression. The Paratethys Sea plooded different terrains in Fig. 3. Stratigraphic distribution of selachian occurrences across the southern mar Serbia that were completely gin of the Pannonian Basin Systems. Stratigraphic correlation table and position of emerged or covered with la- the studied interval. Global chronostratigraphy after HILGEN et al. (2009), magne custrine deposits during ear- tostratigraphy and calcareous nannoplankton zones from LOURENS et al. (2004a, b), ly Miocene. PILLER et al. (2007), HOHENEGGER et al.(2009), PEZELJ et al. (2013), SANT et al. (2017). Consistent to other geo- The position of calcareous nannoplankton in the study area after KRSTIĆ (1978), graphically close-by localities MIHAJLOVIĆ & K NEŽEVIĆ (1989), JOVANOVIĆ (2018), modiFied. (S CHULTZ, 2013; SZABÓ & KOCSIS, 2016; TRIF et al., 2016; TRIF & CODREA, 2017), the observed BIGNY, 1846). However, the benthic foraminifera sharks appeared to be common in the Badenian de- Frondicularia sculpta (K ARRER, 1862) and Annulo posits all around the Central Paratethys (SZABÓ & KOC- frondicularia annularis (D’ORBIGNY, 1846) from ‘Višn- SIS, 2016) (Fig.4). In comparison with other localities jica Clay’ (GRUJIČIĆ, 2010), Austria (FLÜGEL, 1960) and in Croatia, Bosnia and Serbia (southern margin of Romania (POPESCU & CRIHAN, 2004) are indicative for Pannonian Basin Systems of Central Paratethys), Early Badenian (Lagenidae Zone). Based on nanno- Višnjica is richer in shark remains. In addition to the

34 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

Fig. 4. Regional distribution of shark fauna; Otodus (Megaselachus) megalodon Agassiz, 1835 (white circles), Hemipristis serra Agassiz, 1835 (gray circles), and Odontaspididae Müller and Henle, 1839 (black circles) in Middle Miocene (Baden ian), sensu SZABÓ & K OCSIS (2016) in the Central Paratethys, modiFied.

taxa herein reported, Grubić (1958) mentioned from Transition. The preserved Pinus cones and carbonated Višnjica Carcharodon megalodon, Lamna cuspidata fossil wood fragments, as well as the analysis of the and Lamna (Otodus) appendiculata. In northern Cro- molluscan fauna indicate subtropical marine waters, atia sharks are found in Papuk Mountain, but proba- with depths exceeding sometimes 100 m. The Baden- bly reworked in Sarmatian sediments (VRSALJKO et al., ian VCF was deposited in a neritic environment and 2010). The dominant shark family from Višnjica is the Aturia shell accumulations at Višnjica were inter- the Odontaspididae documented by seven teeth preted by STEVANOVIĆ (1970) as post mortal processes, which could be identipied only at a family level. The the shells having drifted into shallower marine envi- rarity of shark teeth in the Badenian of Serbia could ronments. Unlike Višnjica, some areas were covered be linked either to the lack of collecting or to the ab- by the sea only at the end of the Badenian, or in the sence of equivalent ecological niches. Sarmatian. Several outcrops have shown that the The Višnjica Clay, besides foraminifera, gastropods transgression had occurred in Early Badenian on the and bivalves, also bears shells of scaphopods, carbon- basis of the plooding areas located more than 200 km ated fossil wood, three-dimensional preserved Pinus southern of Belgrade (Koceljeva, Aranđelovac) (AN- cones and pyritized shells of Aturia aturi (BASTEROT, ĐELKOVIĆ et al., 1989). However, in such areas, the sea 1825). The fossil molluscan assemblages indicate a was mostly very shallow. Tectonic movements sepa- relatively global warm period (Middle Miocene Cli- rated these rocks into distinct blocks, lifted or matic Optimum), before the Middle Miocene Climatic plunged, during the Neoalpine tectonic events

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 35 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

(MAROVIĆ et al., 2007, TOLJIĆ et al., 2014). For this rea- Conclusion son Badenian sediments are rarely exposed at the surface. The common presence of some large, oceanic The shark teeth from VCF document three diffe- sharks (e.g., Otodus, Cosmopolitodus) in the Central rent families: Otodontidae, Hemigaleidae and Odon- Paratethys during Middle Miocene could be explained taspididae, described for the pirst time from the by the abundance of their potential prey, represented Badenian of Serbia. The pirst two families are docu- by diverse marine mammals such as whales and dol- mented by a single species, i.e. Otodus (Megasela phins (SZABÓ & KOCSIS, 2016). Some remains of Ce chus) megalodon Agassiz and Hemipristis serra totherium cf. rathkii Brandt, 1843 were also found in AGASSIZ. The shark assemblage includes well known the Badenian deposits of northern Bosnia (STEFANOVIĆ, and widespreaded Middle Miocene (Badenian) taxa, 2010). Compared to the recent similar top predator reported also from the other Paratethyan localities. Carcharodon carcharias, habitat of O. (M.) megalodon Therefore, the Middle Miocene of Serbia is domi- was probably similar. This shark hunted both on nated by the cosmopolitan sharks. We conclude that coastal environments but also offshore, on the conti- the shark pindings are consistent with a connection nental and island shelves, over hundreds of meters between the southern margin of the Pannonian Basin (WENG et al., 2007). Systems and the other basins and sub-basins within While O. (М.) megalodon was clearly the top pre- the Central Paratethys in the Badenian. Višnjica rep- dator in these environments, we consider H. serra, resents the richest Badenian shark teeth locality in with a length of 4 to 5 m (KENT, 1994) to be similar Serbia. The lack of small shark teeth is most probably in its trophic level. The actual representative of the related to the sampling due to the fact that all the genus, H. elongata KLUNZINGER 1871, has a very simi- shark teeth were obtained by picking from the ex- lar dentition, but the smaller teeth length. Following posed sediment surface without sieving. COMPAGNO (1984), the recent H. elongata feeds with a variety of pish prey, including smaller sharks and rays. Presumably, the fossil H. serra had a similar diet. Acknowledgments The recent Hemipristis elongata lives in depths of up to 132 m (STEVENS & MC LOUGHLIN, 1991). We thank both reviewers, for their suggestions and Many Badenian localities have yielded a high diver- comments on our manuscript: Werner Schwarzhans (Na- sity and excellent preservation of fossils, mostly inver- tural History Museum of Denmark, Zoological Museum, tebrates. The shark teeth from Višnjica however Denmark) and Vasja Mikuž (University of Ljubljana, Depart- display different colors and marks of damage that ment of Geology, Slovenia). Also, the authors thank Oleg could be explained by their specipic taphonomy. All the Mandić (Natural History Museum, Vienna) for his observa- shark teeth are considered autochthonous. As the tions and suggestions for the stratigraphic interpretation. Badenian transgression overlies continental and plu- One of the authors (VC) would also like to acknowledge the vio-lacustrine older deposits, reworking was not pos- support received from the Babeș-Bolyai University, Cluj- sible. Nowadays, the Višnjica exposure is inaccessible Napoca, grants AGC: 30852, 30851, 30850, 31509. any more. A single tooth of Otodus (M.) megalodon is white in color. This specipic color leads us to think that it originated from a different, distinct layer, possibly References the white Leitha-type sandstone (once, exposed at Bela Stena, near Višnjica). The enamel and roots of the AGASSIZ, L. 1833-1843. Recherches sur les poissons fossiles, Odontaspididae teeth is heavily worn, most probably (Contenant l’Histoire des Cycloides). Imprimerie de Pe- due to the wave action. Small transport could damage titpierre, Neuchatel. Suise, 390 pp. the root of the teeth. Even moderate abrasion/corro- ANĐELKOVIĆ, M., PAVLOVIĆ, M., EREMIJA, M. & ANĐELKOVIĆ, J. 1989. sion will result in reshaping of the edges of the root Paleogeograpija. In: ANĐELKOVIĆ, M., (Ed.). Geologija šire and a rounding of the lingual protuberance in basal okoline Beograda, [Geology of the surroundings of Bel- and propile views (SIVERSSON et al., 2015). grade - in Serbian], IV, University of Belgrade, 282 pp.

36 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

BERG, L.S. 1958. System der Rezenten und Fossilen Fischartigen ĐURIĆ, D., BOGIĆEVIĆ, K., PETROVIĆ, D. & NENADIĆ, D. 2017. Late und Fische. Hochschulbücher für Biologie, Berlin, 310 pp. Pleistocene Squamate Reptiles from the Baranica Cave BRANDT, J.F. 1843. De Cetotherio, novo balaenarum familiae near Knjaževac (Eastern Serbia). Geološki anali Bal genere, in Rossia meridionali anti aliquot annos eff oso. kanskoga poluostrva, 78: 23–35. Bulletin de l’Academie imperiale des Sciences de St. Pe FLÜGEL, E. 1960. Typen-Katalog. Verzeichnis der in der Geo- tersbourg, (2) 1: 145–148. logisch-Paläontologischen bteilung des Naturhistori- BRZOBOHATÝ, R. & SCHULTZ, O. 1978. Die Fischfauna des Ba- schen Museums in Wien aupbewahrten Typen sowie deniens. In: PAPP, A., CICHA, I., SENES, J. & STEININGER, F., der bbildungsoriginale I. Invertebrata: 1. Protozoa. 2. (Eds.). M4 Badenien (Moravien, Wielicien, Kosovien). Coelenterata. Geologie und Paläontologie. Annalen des Chronostratigraphie und Neostratotypen, Miozän der Naturhistorischen Museums in Wien, 64: 65–104. Zentralen Paratethys, 6: 441–465. FLÜGEL, E. & KOLLMANN, H. 1964. Verzeichnis der wichtigsten CAPPETTA, H. 2012. Handbook of Paleoichthyology. Vol. 3E: Objekte der Geologisch-Paläontologischen Sammlung Chondrichthyes. Mesozoic and Cenozoic Elasmobranchii: (Rotpunkt-Verzeichnis). Alphabetische Zusammenstel- Teeth. Verlag Dr. Friedrich Pfeil. München, 512 pp. lung wichtiger Schauobjekte mit Erläuterungen. Veröf CARRILLO-BRICENO, J.D., DE GRACIA, C., PIMIENTO, C., AGUILERA, fentlichungen aus dem Naturhistorischen Museum Wien, O.A.,KINDLIMANN, R., SANTAMARINA, P. & JARAMILLO, C. 2015. 5: 148–156. A new Late Miocene Chondrichthyan assemblage from FOWLER, H.W. & KUMMEL, H.B. 1911. A Description of the the Chagres Formation, Panama. Journal of South Amer Fossil Fish Remains of the Cretaceous, Eocene and ican Earth Sciences, 60: 56–70. Miocene Formations of New Jersey. Bulletin of New Jer CHIRA, C. & MĂRUNŢEANU, M. 1999. Middle Miocene (Lower sey Geological Survey, 4: 1–192. Badenian) calcareous nannofossils from the Mureş GLICKMAN, L .S. 1964. Sharks of Paleogene and their strati passage way and Făget basin, Romania. Acta Palaeon graphic signiFicance. Nauka Press Moscow, 228 pp. tologica Romaniae, 2: 73–82. GRUBIĆ, D. 1958. Prilog poznavanju Višnjičkih plavih glina CIONE, A.L., MENNUCCI, J., SANTALUCITA, F. & ACOSTA HOSPITALE- [Contributtion to knowledge of Višnjica blue clays – in CHE, C. 2007. Local extinction of genus Carcharias (Elas- Serbian]. Vesnik Zavoda za geološka i geoFizička istraži mobranchii, Odontaspididae) in the eastern Pacipic vanja N.R. Srbije, 15: 11–121. Ocean. Revista Geológica de Chile, 34: 139–145. GRUJIČIĆ, LJ. 2010. Foraminifere roda Frondicularia iz COMPAGNO, L.J.V., 1973. Interrelationships of living elasmo- Beogradskog dunavskog ključa [Foraminifers of the branchs. Zoological Journal of the Linnean Society, 53 Genus Frondicularia from Belgrade area Danubian Me- (Supplement 1): 15–61. ander]. In: BANJAC, N. (Ed.),. Proceedings of the 15th Con- COMPAGNO, L.J.V. 1984. An annotated and illustrated cata- gress of the Geologists of Serbia with international logue of shark species known to date (Carcharini- participation, Belgrade, 69–73. formes). FAO Species Catalogue for Fishery Purposes. HASSE, J.C.F. 1879-1885. Das Natürliche System der Elasmo- Food and agriculture organization of the United Na branchier auf Grundlage des Baues und der Entwicklung tions, 4 (2): 251–655. ihrer Wirbelsäule. Eine Morphologische und Paläonto- ĆORIĆ, S., PAVELIĆ, D., RÖGL, F., MANDIC, O., VRABAC, S., AVANIĆ, logische Studie. Gustav Fischer Verlag, Jena. Allgemeiner R., JERKOVIĆ, L., VRANJKOVIĆ, A. 2009. Revised Middle Theil: i-vi, 1–76, 1879; Besonderer Theil, (1): 1-94, pls. Miocene datum for the initial marine plooding of North 1–12: 1882; (2): 97-109, pls. 13-23: 1882; (3): i-vi, 183– Croatian basin (Pannonian Basin System, Central 285, pls. 24–40: 1882; Ergänzungsheft, pp: 1–27. Paratethys). Geologica Croatica, 62 (1): 31–43. HIDEN, H.R., 1995. Elasmobranchier (Pisces, Chondricht- DE MUIZON, C. &DE VRIES, T.H. 1985. Geology and paleontol- hyes) aus dem Badenium (Mittleres Miozän) des Stei- ogy of late Cenozoic marine deposits in the Sacaco area rischen Beckens (Österreich). Mitteilungen der (Peru). Geologische Rundschau, 74 (3): 547–563. Abteilung für Geologie und Paläontologie am Landes DOMNING, D.P. &PERVESLER, P. 2012. The sirenian Metaxy museum Joanneum, 52/53: 41–110. therium (Mammalia: Dugongidae) in the Badenian HILGEN, F.J., ABELS, H.A., IACCARINO, S., KRIJGSMAN, W., RAFFI, I., (Middle Miocene) of Central Europe. Austrian Journal SPROVIERI, R., TURCO, E. & ZACHARIASSE, W.J. 2009. The of Earth Sciences, 105 (3): 125–160. Global Stratotype Section and Point (GSSP) of the Ser-

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 37 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

ravallian Stage (Middle Miocene). Episodes, 32: KENT, B.W., 1994. Fossil Sharks of the Chesapeake Bay Re 152–166. gion. Egan Rees and Boyer, Columbia, 146 pp. HOHENEGGER, J., RÖGL, F., ĆORIĆ, S., PERVESLER, P., LIRER, F., RO- KLUNZINGER, C.B. 1871. Synopsis der Fische des Rothen ETZEL, R., SCHOLGER, R. & STINGL, K. 2009. The Styrian Meeres II. Theil. Verhandlungen der Königlischen Zoo Basin: A key to the Middle Miocene (Badenian/Lan- logischen–Botanischen Gesellschaft in Wien, 21: ghian) Central Paratethys transgressions. Austrian 441–688. Journal of Earth Science, 102: 102–132. KOCH, A. 1903. Tarnócz, in Nógrád County, as new and rich HOHENEGGER, J., ĆORIĆ, S. & WAGREICH, M. 2014. Timing of the locality for fossil shark teeth. Földtani Közlöny, 33: 22– Middle Miocene Badenian Stage of the Central 44. Paratethys. Geologica Carpatica, 65 (1): 55–66. KOLLMANN, H. 1969. Verzeichnis der wichtigsten Objekte HOLEC, P. & SABOL, M., 1996. The Tertiary Vertebrates from der Geologisch-Paläontologischen Sammlung (Rot- Devínska Kobyla. Mineralia Slovaca, 28: 519–522. punkt-Verzeichnis). Alphabetische Zusammenstellung HOLEC, P. 2001. Chondrichthyes and Osteichthyes (Verte- wichtiger Schauobjekte mit Erläuterungen. Veröffent brata) from Miocene of Vienna Basin near Bratislava lichungen aus dem Naturhistorischen Museum Wien, 5 (Slovakia). Mineralia Slovaca, 33 (2): 111–134. (2): 167–177. IVKOVIĆ, A, VUKOVIĆ, A, NIKOLIĆ J, KOVAĆEVIĆ, D, PALAVESTRIĆ, LJ., KOVÁČ, M., ANDREYEVA-GRIGOROVICH, A., BAJRAKTAREVIĆ, Z., BRZO- PETROVIĆ, LJ., JOVANOVIĆ, O., TRIFUNOVIĆ, I. & SEBINOVIĆ, LJ. BOHATÝ, R., FILIPESCU, S., FODOR, L., HARZHAUSER, M., NAGY- 1966. Osnovna Geološka karta Jugoslavije 1:100,000, MAROSY, A., OSZCZYPKO, N., PAVELIĆ, D., RÖGL, F., SAFTIĆ, B., list Pančevo [Basic Geologic Map of Former Yugoslavia SLIVA, L. & STUDENCKA, B. 2007. Badenian evolution of 1:100,000, Sheet Pančevo]. Savezni geološki zavod, the Central Paratethys Sea: paleogeography, climate Beograd. and eustatic sea-level changes. Geologica Carpathica, JORDAN, D.S. & HANNIBAL, H. 1923. Fossil sharks and rays of 58 (6): 579–606. the Pacipic Slope of North America. Bulletin of the KRSTIĆ, N. 1978. Miocenski ostrakodi iz Dunavskog ključa Southern California Academy of Sciences, 22: 27–63. kod Beograda [Miocene ostracods from Belgrade area JOVANOVIĆ, G. 2018. Badenske školjke jugoistočnog oboda Danubian Meander – in Serbian]. Proceedings of the 9th panonskog basena [Badenian bivalves of the south- Congress of the Geologists of Yugoslavia, Sarajevo, eastern margin of Panonian Basin Systems (Central 117–125. Paratethys) – in Serbian]. Bulletin of Natural History KRSTIĆ, N., SAVIĆ, LJ. & JOVANOVIĆ, G. 2012. The Neogene Lakes Museum, Special editions 44: 1–136. on the Balkan Land. Geološki anali Balkanskoga poluos JOVANOVIĆ, G. & BOŠNJAK, M. 2016. Fissidentalium badense trva, 73: 37–60. (Partsch in Hörnes, 1856) from the Badenian deposits LASKAREV, V. 1924. Sur les equivalents du Sarmatien supé- of the south and southwestern margin of the Pannon- rieur en Serbie. In: VUJEVIĆ, P. (Ed.): Receuil de traveaux ian Basin System (Central Paratethys). Geologica Croat offert à M. Jovan Cvijic par ses amis et collaborateurs. ica, 69 (2): 195–200. Državna Štamparija, Beograd, 73–85. JOVANOVIĆ, G. & TOMIĆ, Z. 1997. Paleoekološke i mineraloške LOURENS, L.J., HILGEN F.J., LASKAR, J., SHACKLETON, N.J. & WILSON, metode pri rekonstrukcijama životnih uslova u baden- D. 2004a. The Neogene period. In: GRADSTEIN, F.M., OGG, skom veku Golupca [Paleoecological and mineralogi- J.G. & SMITH, A.G. (Eds.). A geologic time scale 2004. cal methods in living conditions reconstruction in Cambridge University Press, 409–440. Badenian age at Golubac (Eastern Serbia) – in Ser- LOURENS, L., HILGEN, F., SHACKLETON, N.J., LASKAR, J. & WILSON, bian]. Geološki anali Balkanskoga poluostrva, 61 (2): D. 2004b. Orbital tuning calibrations and conversions 371–392. for the Neogene Period. In: GRADSTEIN, F.M., OGG, J.G. & JOVANOVIĆ, G., ĆORIĆ, S. & VRABAC, S. 2018. Prvi dokazi donjo- SMITH, A.G., (Eds.). A Geologic Time Scale 2004. Cam- badenske morske transgresije kod Koceljeve [The First bridge University Press, Cambridge, 469–484. evidence of marine Badenian transgression near Ko- LUKOVIĆ, M. 1922. Drugomediternakse facije iz okoline celjeva (western Serbia) – in Serbian ]. Abstract Pro Beograda [Second Mediterranean facies from Belgrade ceedings of 17th Serbian Geological Congress, May surroundings – in Serbian].Geološki anali Balkanskoga 17th20th, Vrnjačka Banja, Serbia, volume 1, 139–143. poluostrva, 7 (1): 22–41.

38 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

MAROVIĆ, M., TOLJIĆ, M., RUNDIĆ, LJ. & MILIVOJEVIĆ, J., 2007. navski ključ” – in Serbian]. Geološki anali Balkanskoga Neoalpine tectonics of Serbia. Serbian Geological Soci- poluostrva, 7 (1): 1–21. ety, Belgrade, 87 pp. PAWLOWSKA, K. 1960. The pish remains of the Miocene lime- MIHAJLOVIĆ, Đ. & KNEŽEVIĆ, S. 1989. Krečnjački nanoplankton stone of Pinczowa. Acta Palaeontologica Polonica, 5 (4): iz badenskih i sarmatskih sedimenata Višnjice i Ka - 421–432. raburme kod Beograda [Calcareous nannoplankton PETROVIĆ, M. 1970. Kredni i miocenski foraminiferi Beo- from Badenian and Sarmatian deposits at Višnjica and grada i njegove okoline [Cretaceous and Miocene fora- Karaburma in Belgrade – in Serban]. Geološki anali minifera of Belgrade area – in Serbian]. Geološki anali Balkanskoga poluostrva, 53: 342–366. Balkanskoga poluostrva, 35: 191–209. MIKUŽ, V. 2000. The great-teeth shark Carcharocles mega PETROVIĆ, M. 1985. Foraminiferske zone i loze badenskog lodon (Agassiz) also from Middle Miocene-Badenian kata okoline Beograda [Foraminiferal zone and li- beds above Trbovlje, Slovenia. Geologija, 42: 141–150. neages of the Badenian stage of the surroundings of MIKUŽ, V. & ŠOSTER, A. 2013. A Mackerel Shark (Mega Belgrade - in Serbian]. Geološki anali Balkanskoga po selachus megalodon) pind in Orehovica, Dolenjska, luostrva, 49: 271– 291. Slovenia. Folia Biologica et Geologica, 54 (1): 109–119. PILLER, W.E., HARZHAUSER, M. & MANDIC, O. 2007. Miocene MIKUŽ, V., ŠOSTER, A. & ULAGA, S. 2014. Megaselahus in the Central Paratethys stratigraphy – current status and Miocene beds of Retje – Plesko Quarries above Tr- future directions. Stratigraphy, 4 (2/3): 151–168. bovlje, Slovenia. Folia Biologica et Geologica, 55 (1): PIMIENTO, C., EHRET, D.J., MACFADDEN, B.J. & HUBBELL, G. 2010. 73–88. Ancient Nursery Area for the Extinct Giant Shark MIKUŽ, V., ŠOSTER, A., STARE, F. & SUKIČ-PREKMURSKI, M. 2015. Megalodon from the Miocene of Panama. PLoS ONE, 5 Megalodon teeth from Miocene marlstone at Virštanj, (5): e10552. Slovenia. Folia Biologica et Geologica, 56 (2): 77–107. PIMIENTO, C., GONZALEZ-BARBA, G., EHRET, D.J., AUSTIN, J.W., MAC- MIKUŽ, V., ŠOSTER, A. & ULAGA, Š. 2017. Fossil pish teeth from FADDEN, B.J. & JARAMILLO, C. 2013. Sharks and rays sites between Trbovlje and Laško, Slovenia. Folia Bio (Chondrichthyes, Elasmobranchii) from the late logica et Geologica, 58 (1): 59–75. Miocene Gatun formation of Panama. Journal of Pale MIKINČIĆ, V. 1932. Kenozojski sedimenti između Golupca, ontology, 87 (5): 755–774. Vukotića i V. Gradišta [Cenozoic deposits between Gol- PEZELJ, D.J., MANDIĆ, O. & ĆORIĆ, S. 2013. Paleoenvironmental ubac, Vukotić and V. Gradište – in Serbian]. Bulletin of dynamics in the southern Pannonian Basin duing ini- the Yugoslavian Royal Geological Institute, 1 (1): 89–104. tial Middle Miocene marine plooding. Geologica Car MÜLLER, J. & HENLE, F.G.J. 1838. On the generic characters pathica, 64 (1): 81–100. of cartilaginous pishes, with descriptions of new gen- POPESCU, G. & CRIHAN, I.M. 2004. Contributions to the knowl- era. Magazine of natural history and journal of zoology, edge of Miocene foraminifera from Romania: Super- botany, mineralogy, geology and meteorology, 2: 33-37, family Nodosariacea (Fam. Nodosariidae and 88–91. Vaginulinidae). Acta Paleontologica Romaniae, 4: MÜLLER, A. 1999. Ichthyofaunen aus dem atlantischen Ter 385–402. tiär der USA. Leipziger Geowissenschaften, 9/10: POPOV, S.V., RÖGL, F., ROZANOV, A.Y., STEININGER, F. F., SHCHERBA, 1–360. I.G. & KOVÁC, M. 2004. Lithological paleogeographic PAUNOVIĆ, M. 1987. Ein Beitrag zur Kenntnis der obermio- maps of Paratethys. 10 Maps Late Eocene to Pliocene. zänen Knorpelpische Kroatiens. Jugoslavenske akade Courier Forschungs Institut Senckenberg, 50: 1–46. mije znanosti i umjetnosti, 431: 131–140. PROBST J. 1878. Beiträge zur Kenntniss der fossilen Fische PAVLOVIĆ, P. 1897. Prilog poznavanju foraminifera iz dru- aus der Molasse von Baltringen. Haypische. Jahreshefte gomediternskih slojeva u Srbiji [Contribution to the des Vereins für vaterländische Naturkunde in Württem knowledge of foraminifera from the II Mediterranean berg, 34: 113–154. layers in Serbia – in Serbian]. Bulletin of Serbian Royal PURDY, R. W., SCHNEIDER, V. P., APPLEGATE, S. P., MCLELLAN, J. H., Academy, 56: 114–140. MEYER, R. L. & SLAUGHTER, R. 2001. The Neogene sharks, PAVLOVIĆ, P. 1922. Geološki sastav Beogradskog Dunavskog rays, and bony pishes from Lee Creek Mine, Aurora, ključa [Geological composition of the “Beogradski Du- North Carolina. In: Ray, C. & Bohaska, D.(Eds.). Geology

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 39 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

and paleontology of the Lee Creek Mine, North Carolina, Republic, Vienna Basin. Annalen des Naturhistorischen III, Smithsonian Contributions to Paleobiology, 90: Museums in Wien, 112 (A): 489–506. 71–202. SIVERSSON, M., LINDGREN, J., NEWBREY, M.G., CEDERSTRÖM, P. & RADOVIĆ, P. & BRADIĆ-MILINKOVIĆ, K., 2018. A new elephan- COOK, T.D. 2015. Cenomanian–Campanian (Late Creta- toid dental specimen from the Miocene of Kruševac ceous) mid-palaeolatitude sharks of Cretalamna ap- Basin in Central Serbia. Geološki anali Balkanskoga pendiculata type. Acta Palaeontologica Polonica, 60 poluostrva, 79 (2): 1–10. (2): 339–384. RADWANSKI, A. 1965. A contribution to the knowledge of STEFANOVIĆ, I. 2010. Note on the pirst fossil remains of a Miocene Elasmobranchii from Pinczow (Poland). Acta whale from northern Bosnia. Geološki anali Balkan Palaeontologica Polonica, 10 (2): 267–279. skoga poluostrva, 71: 127–137. RÖGL, F. 1998. Palaeogeographic Considerations for Me- STEVANOVIĆ, P.1970. Paleogeografsko-ekološke prinove iz tor- diterranean and Paratethys Seaways (Oligocene to tona okoline Beograda [Paleogeographic-ecological nov- Miocene). Annalen des Naturhistorischen Museums in elties from the Tortonian of Belgrade area– in Serbian]. Wien, 99 (A): 279–310. Bulletin of Serbian Academy of Science and Arts, 278, De RÖGL, F. & STEININGER, F.F. 1984. Neogene Paratethys, partment of Natural Science and Mathematics, 33: 1–20. Mediterranean and Indo-pacipic Seaways. Implications STEVANOVIĆ, P. 1977. Miocene of Belgrade. In: PETKOVIĆ, K. for the paleobiogeography of marine and terrestrial (Ed.). Geology of Serbia, 2-3, Stratigraphy of Cenozoic, biotas. In: BRENCHLEY, P. (Ed.). Fossils and Climate. John 3-8. University of Belgrade, 107–171. Wiley and Sons, 171–201. STEVENS, J.D. & MCLOUGHLIN, K.J. 1991. Distribution, size and RÖGL, F., ĆORIĆ, S., HARZHAUSER, M., JIMENEZ-MORENO, G., KROH, sex composition, reproductive biology and diet of A., SCHULTZ, O., WESSELY, G. & ZORN, I. 2008. The Middle sharks from northern Australia. Australian Journal of Miocene Badenian stratotype at Baden-Sooss (Lower Marine and Freshwater Research, 42 (2): 151–199. Austria). Geologica Carpatica, 59: 367–374. SZABÓ, M. & KOCSIS, L. 2016. A new Middle Miocene sela- SANT, K., PALCU, D., MANDIC, O. & KRIJGSMAN, W. 2017. Chang- chian assemblage (Chondrichthyes, Elasmobranchii) ing seas in the Early–Middle Miocene of Central Eu- from the Central Paratethys (Nyirád, Hungary): impli- rope: a Mediterranean approach to Paratethyan cations for temporal turnover and biogeography. Geo stratigraphy. Terra Nova, 29 (5): 273–281. logica Carpathica, 67 (6): 573–594. SCHULTZ, O. 1971. Die Selachier-Fauna (Pisces, Elasmobran- TOLJIĆ, M., NENADOVIĆ, D., STOJADINOVIĆ, U., GAUDÉNYI, T. & chii) desWiener Beckens und seiner Randgebiete im BOGIĆEVIĆ, K., 2014. Quaternary tectonic and deposi- Badenien (Miozän). Annalen des Naturhistorischen Mu tional evolution of eastern Srem (northwest Serbia). seums in Wien, 75: 311–341. Geološki anali Balkanskoga poluostrva, 75: 43–57. SCHULTZ, O. 1977. Elasmobranch and teleost pish remains TRIF, N., CIOBANU, R. & CODREA, V. 2016. The pirst record of from the Korytnica Clays (Middle Miocene, Holy Cross the giant shark Otodus megalodon (Agassiz, 1835) from Mountains, Poland). Acta Geologica Polonica, 27 (2): Romania. Brukenthal Acta Musei, 11 (3): 507–526. 201–209. TRIF, N. & CODREA, V. 2017. Some Badenian pish teeth from SCHULTZ, O. 1979. Supplementary notes on elasmobranch Western Transylvania (Romania). Oltenia. Studii şi and teleost pish remains from the Korytnica Clays (Mid- comunicări. Ştiinţele Naturii, 33 (1): 7–17. dle Miocene; Holy Cross Mountains, Central Poland). VRSALJKO, D., JAPUNDŽIĆ, S., KOVAČIĆ, M., GRGANIĆ-VRDOLJAK, Z. & Acta Geologica Polonica, 29 (3): 287–293. PLEŠE, P. 2010: Vranić: The most important pinding SCHULTZ, O. 2013. Pisces. Ein systematisches Verzeichnis place of fossil whales in Northern Croatia. 4th Croatian aller aufösterreichischem Gebiet festgestellen Fossilien Geological Congress, Zagreb, 118. Band 1/Teil 3: Bivalvia neogenica (Solenoidea-Clava- WENG, K.C., BOUSTANY, A.M., PYLE, P., ANDERSON, S.D., BROWN, gelloidea). In: PILLER, W. (Ed.). Catalogus Fossilium Aus A. & BLOCK, B.A. 2007. Migration and habitat of white triae, Bd. 3, Verlag der Österreichischen Akademie der sharks (Carcharodon carcharias) in the eastern Pacipic Wissenschaften, Wien, 576 pp. Ocean. Marine Biology, 152 (4): 877–894. SCHULTZ, O., BRZOBOHATÝ, R. & KROUPA, O. 2010. Fish teeth ZAPFE, H., 1954. Vorzeitliche Meere im Wiener Becken. Uni- from the Middle Miocene of Kienberg at Mikulov, Czech versum. Natur und Technik, 9 (15): 468–480.

40 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

Резиме две породице су представљене по једном врстом - Otodus (Megaselachus) megalodon AGASSIZ и Hemi- Средњомиоценски зуби ајкула из pristis serra AGASSIZ. Примерци Odontaspididae, због јужног обода Панонског басена оштећења нису могли бити прецизније одређени. (Србија, Централни Паратетис) Иако је анализиран мали број фосилних остатака, очигледно је да су у средњомиоценским седимен- Миоценски зуби ајкула из Паратетиса су по- тима Србије биле заступљене космополитске знати у геолошким радовима још од 19. века. врсте ајкула. Можемо закључити да су ови таксо- Најновија истраживања су омогућила процену ни доказ о везама јужног обода Панонског басена разноврсности, као и њихову географску и стра- са другим басенима унутар Централног Парате- тиграфску дистрибуцију (SZABÓ & KOCSIS, 2016). На тиса, током бадена. Разноврсност морске фауне подручју јужног обода Панонског басена остаци из Вишњице, као и других баденских локалитета, миоценских зуба ајкула су релативно ретки и указује на миграцију фауне из Медитеранског углавном су представљени врстама које на по- басена, преко Словеначког коридора у област дручју Централног Паратетиса имају широко јужног обода Панонског басена. Везе су констато- географско распрострањење. Описани таксони ване и између Панонског и Трансилванијског попуњавају празнину у познавању средњомио- басена, преко пролаза Муреш (CHIRA, C. & MĂRUNŢE- ценске фауне ајкула у овом региону и омогућа- ANU, 1999). Ова специфична палеогеографија вају корелације са сличним фаунама из других омогућила је заједничке особине не само у фау- локалитета Централног Паратетиса. нама мекушаца (JOVANOVIĆ, 2018), већ и у заједни - Стратиграфија Формације Вишњичких глина цама морских кичмењака. Локалитет Вишњица (VCF) је предмет расправа геолога, и сви распо - је најбогатије налазиште зуба ајкула у Србији, ложиви подаци указују на средњомиоценску иако је фосилна асоцијација малобројна. Зуби из старост (баден, кречњачки нанопланктон NN5 Вишњице су различите боје, а запажена су и ра- зона). Чести налази остатака зуба прилично ве- зличита оштећења, што би могло бити објашње- ликих ајкула (нпр. Otodus, Cosmopolitodus) у сред- но специфичном тафономијом. Док су налази њем миоцену Централног Паратетиса могли би ситних Elasmobrachii у другим средњомиоцен - бити објашњени обиљем њиховог потенци јалног ским налазиштима Паратетиса чести (SZABÓ & плена, представљеног разним сисарима као што KOCSIS, 2016), у српским локалитетима ови остаци су китови и делфини. Станиште предатора Otodus нису пронађени, што не значи да нису били део (Megaselаchus) megalodon као и Hemipristis serra је баденских заједница, већ да се у досадашњим била приобална морска средина са дубинама које истраживањима највероватније није посветила су могле бити нешто веће од 100 м, на шта указују пажња просејавању веће количине седимената, и заједнице других фосилних организама. што би управо требало да буде циљ будућих У овом раду су први пут описани зуби ајкула из теренских радова у Србији. баденских седимената VCF код Београда (Србија), који припадају трима различитим породицама: Manuscript recieved February 10, 2019 Otodontidae, Hemigaleidae и Odontaspididae. Прве Revised manuscript accepted June 25, 2019

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 41 GORDANA JOVANOVIĆ, N ICOLAE TRIF, V LAD CODREA & D RAGANA ĐURIĆ

Plate 1.

Selachians from Višnjica, Badenian (NHMB): ad - col. Petar Stevanović – PS 23; eg - Petar Pavlović – K 7050); hj: Otodus (Megaselachus) megalodon (col. Petar Stevanović – PS 24); kn: Hemipristis serra (col. Petar Stevanović – PS 25); or: Odontaspididae indet; a, e, h, k, o (col. Velimir Milošević – KV-P 282); (lingual view; b, f, j, l, p - labial view; g, h, n - mesial view; d, n - distal view; c - detail of serration).

42 Geol. an. Balk. poluos., 2019, 80 (1), 29–43 Middle Miocene shark teeth from the southern margin of the Pannonian Basin System (Serbia, Central Paratethys)

Geol. an. Balk. poluos., 2019, 80 (1), 29–43 43