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

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; ([email protected]) 2 Croatian Natural History Museum, Department of Geology and Paleontology, Demetrova 1, Zagreb, Croatia; (corresponding author: [email protected]) doi: 10.4154/gc.2016.20

Abstract Article history: The Badenian (Middle Miocene) Scaphopoda from the south and southwestern margin of the Manuscript received December 16, 2015 Pannonian Basin System (Central Paratethys) are poorly documented, and in published papers Revised manuscript accepted May 31, 2016 mostly presented by short lists of identified species. For the present study, 300 scaphopod speci­ Available online June 28, 2016 mens were examined with the focus on a singular species, 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 Keywords: Fissidentalium badense, Badenian, Belgrade (Serbia), while the others originate from the vicinity of Zagreb (Croatia) and Ugljevik Pannonian Basin System, Central Paratethys (Bosnia and Herzegovina).

1. INTRODUCTION from the investigated area belong palaeogeographically to the The Scaphopoda is a class of , also known as the “tusk south and southwestern margin of the Central Paratethys, and shells”. Their shell is tubular and generally curved, having open­ geotectonically to the Pannonian Basin System, surrounded by ings at both ends. Scaphopods are a cosmopolitan marine infau­ the Alps, Carpathians and Dinarides (Fig. 1) (e.g. RÖGL 1998, nal species, and live from the intertidal zones down to depths of 1999; PAVELIĆ, 2001, 2002; PILLER et al., 2007). more than 6000 m. Scaphopod species mostly burrow in sand or The beginning of the Badenian epoch is marked by a marine mud, with their head pointed downwards, and the narrow poste­ transgression, which flooded the areas behind the newly uplifted rior end of the shell protruding above the substratum (KNUD­ mountain chains (Paratethys) and was also prominent in the Medi­ SEN, 1964; LAMPRELL & HEALY, 1998 and references therein; terranean (Fig. 1) (RÖGL, 1998, 1999; KOVÁČ et al., 2007). Ac­ HABDIJA et al., 2011). cording to numerous papers, the Badenian deposits of the south Although the scaphopod systematics have been revised and southwestern margin of the Central Paratethys lie discorda­ over a few decades, the time of their appearance is still a matter ntly on the older Miocene, Mesozoic or even Palaeozoic deposits. of debate, as are their phylogenetics, so they are considered to be The contact with basal rocks is in many places unclear or erosive a puzzling molluscan class. Two orders of scaphopods are or tectonic in character (e.g. ŠIKIĆ et al., 1978, 1979; BASCH, known: and Gadilida, which differ both morphologi­ 1983a, b; VRABAC & MIHAJLOVIĆ, 1990; ANĐEL­KOVIĆ et cally, in shell and foot form, and behaviourally, in different bur­ al., 1991; PAVELIĆ, 2001, 2002; SAVIĆ et al., 2005; PIKIJA, rowing activities (LAMPRELL & HEALY, 1998 and references 2009). therein; STEINER & KABAT, 2001, 2004). At the southern margin of Central Paratethys, F. badense is This paper focuses on the Badenian dentaliid species Fissi­ recorded in the Lower Badenian deposits of Serbia (wider Bel­ badense (PARTSCH in HÖRNES), recorded in the grade region) (PETKOVIĆ et al., 1976), in the assumed Lower to Mio­cene deposits of the Paratethys, and widely distributed in Ba­ Middle Badenian of western Serbia (ŽUJOVIĆ, 1889; PAVLOVIĆ, denian sediments of the Central Paratethys (e.g. BALDI, 1960; 1903; STEVANOVIĆ, 1949; PETROVIĆ, 1967), and in the Lower KOJUMDGIEVA & STRACHIMIROV, 1960; BALLUK, 1972; Badenian (Upper Lagenidae zone) in northeastern Bosnia and TITA, 2007; HARZHAUSER et al., 2011). Specimens of F. ba­ Herzegovina (Bogutovo Selo) (VRABAC et al., 2013, 2014; PE­ dense included here were collected from the Badenian sediments ZELJ et al., 2013) (Fig. 1). In the area of the southwestern margin of the south and southwestern margin of the Central Paratethys of the Central Paratethys, this species is present in the Miocene (Croatia, Bosnia and Herzegovina and Serbia). The analyzed spe­ (probably Lower Badenian) marls of the wider Zagreb region cimens are housed at the Natural History Museum Belgrade (Med­vednica Mt., Čučerje area) (KOCHANSKY, 1944, 1957; (NHMB) and Croatian Natural History Museum (CNHM). The ĆORIĆ et al., 2009), and in the assumed Middle to Upper Bade­ first finding ofF. badense from Bosnia and Herzegovina is pub­ nian deposits of the Samobor area (Vrhovčak locality, previously lished here. known as „Zaprešić Breg“ or „Zaprešić brijeg“) (ŠUKLJE, 1929; PAVLOVSKY, 1957, 1960; BAJRAKTAREVIĆ, 1978; VRSALJ­KO, 2. GEOLOGICAL SETTING 2003; BAKRAČ et al., 2010). In the Badenian sediments of the The Central Paratethys extended from Bavaria to the Carpathian Glinsko Pokuplje area, this species is found at one locality (PI­ mountain chain (e.g. RÖGL 1998, 1999). Badenian sediments LAR, 1873). Geologia Croatica et al., 2012). ning,apical− Hpheight, apicalWp− width (after STEINER, 1999;SILVA-FILHO curvature,height−Ha theventralof opening, Wawidth−theventralof ope re maximum, Larc − maximum length from the apex to the deepestdiameter, point ofDmax the– maximum distance from the anterior opening, Arc – Figurecurvatu 2. Dvorska valley“ collection (3 specimens). Zrinjsko and area Pokuplje Glinsko the from fauna Pliocene and „Miocene of the part are locality third the from Specimens offauna Zaprešić Breg Samobor“ near collection (5 specimens). „Miocene the and (10specimens) collection Mt.“ Medvednica are part of the Miocene „Marine fauna of the southern part of the (1 specimen). mens), and Bogutovo Selospeci (3 Serbia near western Ugljevik, in area Bosnia Loznica the and from Herzegovina come imens nović collection, assembled in 1962 (40 specimens). spec Other P. the Ste and 1897specimens) (90 in assembled collection, P. Pavlovićthe of part a are they and Belgrade),of area (wider shell fragments. as preserved are specimens of majority the and area, apex the mostly in damaged, are specimens analyzed HÖRNES). the All species the to belong Three hundred scaphopod shells were examined, and 152 of them 3. MATERIALS AND METHODS marked localities containing recorded dentaliids Figure 1. 196 CNHM CNHM specimens originating from the wider Zagreb region locality Višnjica the at collected were specimens NHMB Geographic and paleogeographic setting of the investigated area Morphometric measurement parameters: L − length, Max – maximum Fissidentalium badense Fissidentalium

(after KOVÁČ et al., 2007). (PARTSCH in in (PARTSCH

­ with ­va­ ­ ­ - - - 1897 1856 Type species: Subgenus Genus Family Children, 1834 Order Dentaliida da Costa, 1776 Class Scaphopoda Bronn, 1862 SYSTEMATIC PART RESULTS4. (1999) and SILVA 2004). used Parameters in are the STEINER after determination 2001, KABAT CHIMIROV,PAVIA,1960; & 1991; STEINER STRA & KOJUMDGIEVA 1897;SACCO, 1856; HÖRNES, and thereforeserved, chosenhere. to be described NOVIĆ & JOVANOVIĆ, 1998). This fossil fauna is the best pre belonging to them among Višnjica (Fig. 1), with 280 specimens from clayey sediments, 130 flat near the wider end of the shell. Secondary ribs are inserted inserted are ribs Secondary shell. the of end wider the near flat radial ribs are near the apical area. Ribs become lower and almost and moderately 8 Sharp high, to lines. 10 and growth ribs radial solid with ornamented is shell The opening. anterior wider the the collected specimens. The shell narrows and slightly curves to concave, and the ventral is convex. curv Wa = 6.3 Hp mm; = 2.9 Wp mm; = 2.8 mm from Bogutovo Selo near Ugljevik (Bosnia and Herzego 3 specimens from the Glinsko Pokuplje area (Croatia), 1 specimen (Čučerje area), 5 specimens from the „Zaprešić Breg“ locality and Mt.Medvednica the (Serbia), from area specimens 10 damaged cimens from the Višnjica locality, 3 specimens from the Loznica 2013 2011 1998 1991 1960 1960 1897 1947 1929 1925 Determination of species is based upon several papers (e.g. papers severalupon based ofis species Determination is scaphopods of number largest the yielding site The Description Dimensions Material NEROVÁ, p. 20, pl. fig.2, 2 et al.,HARZHAUSER p. 219, pl. 8. figs. 6 JOVANOVIĆ, p. 201–202 PAVIA, p. 146, pl. 5, fig. 4; pl. 6, fig. 6. ( 19–20 KOJUMDGIEVA& STRACHIMIROV, p. 225, pl. 52, figs. BER, p. 302, pl. 7, figs.8–10, pl. 8, fig. 3 KAUTSKY – KAUTSKY, p. 53–54, pl. 5, fig. 12 pl. 9, figs. 17–30 ed, narrow and ­ed, narrow elongated, open at both ends Dentalium () Dentalium badense Entalis badensis Fissidentalium badense Fissidentalium Dentalium Entalis badensis Fissidentalium badense Fissidentalium Dentalium badense Dentalium Dentalium Dentalium badense Dentalium Dentalium Dentalium Dentalium etlu badense Dentalium Fissidentalium badense Fissidentalium Antalis : 20 relatively well preserved and 110 damaged spe­ Dentalium Dentalium ergasticim

( Badense Badense : L = 37.3= L : mm; ( : Shell is medium to large, solid, moderately moderately solid, large, to medium is Shell : ( - Fissidentalium Entalis FILHO et al. (2012) (Fig. 2). Entalis Adams & Adams, 1854 (PARTSCH) – SACCO, p.107, pl. 9, figs. 17–20 (Pl. 1, Figs. 1 Fischer, 1885 PARTSCH (PARTSCH) p. 52 – ŠUKLJE, ) – p.HÖRNES, 652, pl. 50, fig. 30 badensis ) ) F.badense ATCI JVNVĆ & JOVANOVIĆ – PARTSCHI badense (PARTSCH1856) HÖRNES, in – (PARTSCH 1856) HÖRNES, in – (Partsch in Hörnes, 1856) Hörnes, in (Partsch Larc = 32.0 6.4= = Ha mm; Larc mm; ) )

badense (PARTSCH,1856)ROUT – et var. – SACCO, p. 107–108, FISCHER, 1885, monotype PARTSCH in HÖRNES – HÖRNES in PARTSCH The apex is in all damaged ATC var. PARTSCH - – (LUKOVIĆ, 1922; JOVA­ 6) BALDI, p. 57, pl. 1, fig. 5 PARTSCH – TAU – PARTSCH . Geologia Croatica 69/2 Croatica Geologia cum syn.).cum – 7 . Dorsal side is borealis ­vina). ­ ­ ­ ­

Geologia Croatica 197 (PARTSCH in HÖRNES, 1856) fromHÖRNES,Badenianthe in deposits ... (PARTSCH 1856) of ). Fissidentaliumbadense PARTSCH in HÖRNES, 1856 HÖRNES, in PARTSCH ( a (Inv. No. K−2347/1); No. (Inv. a nv. No. K−2689); No. nv. tia (Inv. No. 573); 573); No. (Inv. tia uplje area, Croatia (Inv. No. 1.220). No. (Inv. Croatia area, uplje elo, Bosnia and Herzegovina (Inv. No. K−6746); No. (Inv. Herzegovina and Bosnia elo, „Zaprešić Breg“), Croatia (Inv. No. 1.413); 1.413); No. (Inv. Croatia Breg“), „Zaprešić Fissidentalium badense badense Fissidentalium Vrhovčak ( Vrhovčak Pok Glinsko Višnjica locality, Serbi locality, Višnjica (I Serbia Loznica, S Bogutovo Croa Čučerje,

5 6 1 2 3 4 Plate 1. 1. Plate GordanaMarijaJovanović and Bošnjak: Geologia Croatica GMELIN, GMELIN, dle Badenian, and GRUJIČIĆ (2010) specifiesthe zone with upon foraminifera, STEVANOVIĆ (1977) attributes it to the Mid­ position of the „Višnjica clays“ is not precisely determined. Based which contains many specimens of Baden Pleurothoma clayey sediments, known as the „Višnjica clays“ or the „clays with largest ofnumber specimens. dentaliids, Višnjica near recorded Belgrade (Serbia) the of is the number site yielding the the Given CNHM. and NHMB the at talium badense belong to the dentaliids of Paratethys, the the majority Central of scaphopod discoveries In the Badenian sediments of the south and southwestern margin 5. DISCUSSION MIROV, 1960; PAVIA, 1991; et al., HARZHAUSER 2011). Miocene of the Paratethys (e.g. KOJUMDGIEVA & STRACHI­ Middleof the typical species a represents and distribution, phic 2002; HARZHAUSER et al., 2011). 1897), and in (SACCO, Hills Torinothe the at LowerMediterranean the in MioceneBurdigalian of the Paratethys (HARZHAUSER,lium badense that can contribute to the research of the fossil species. significance and thethe of aperture, numberrib and morphology talium searchof the and ecology of the recent genus 1856) was first recognized as AKOKO, 1999). tionofthescaphopods (e.g. REYNOLDS, 1997; REYNOLDS & face. cross The growth lines (except in the apical area) the characterize shell sur irregular dense and fine ribs, radial Together the width. with in and thickness of the secondary ribs. Concave interrib spaces vary other with only minor differences. The species varies in number posterior side of the shell are almost from indistinguishable each at the ribs shellsecondary and not does exceed 30. primary The of the part posterior the near ribs of inserted number maximum between the ribs primary at various distances from the apex. The 198 HÖRNES), scaphopod specimens: Hungary (BALDI, 1960). The „Višnjica clays“ contain numerous ofSzokoly,zone) sediments (Lagenidae LowerBadenian the in tria (MANDIC & HARZHAUSER, 2003; MANDIC, 2004), and (JOVANOVIĆ,Aus in formation 2014), Gaindorf Grund the in occur in the Lower Badenian of Bukovac on the Fruška gora Mt. also species pectinid These stratum. clay 1922) the VIĆ, cover (SOWERBY) and fine ro subgenusthesingleout Koritnica, Poland (BALLUK, 1972). (1960) has more secondary ribs (35), as do several specimensThespecies from described in KOJUMDGIEVA & STRACHIMIROV locality, and presented here, fits the descriptions by PAVIA DGIEVA(1991). & STRACHIMIROV, 1960).Entalis rutilus carinatus ­rutilus - Remarks: At the VišnjicaAt the the locality, werediscoveredin scaphopods distribution stratigraphic and Geographic shelled pectens ( - (SACCO, 1897) or (FISCHER),HEALY& LAMPRELL (1998) indicatethe Sooss in Vienna (PAVLOVIĆ, Vienna in 1903; Sooss LUKOVIĆ, 1922), Dentalium (Antale) novemcostatum decemcostulata (Antale) novemcostatum Dentalium Gadilinajani “. This site was compared to the Badenian stratotype (PARTSCH in H Many authors studied the phylogeny and classifica­ (PARTSCH in H - section of both isends subcircular. Fissidentalium badense Aequipecten scabrella (d’ORBIGNY). Sandstones containing rare rare containing (d’ORBIGNY). Sandstones Costelamussium . Fissidentalium badense Fissidentalium badense Here, Dentalium Dentalium HÖRNES, HÖRNES, Dentalium badense

we analyzed and revised Ö Ö RNES, RNES, 1856) appears during the RNES, RNES, 1856) specimens stored LINNAEUS 1758.LINNAEUS re the In F. badense F. badense

STEINER & KABAT (2004) ( Entalis F. badense sp.), (PARTSCH in H Dentalium sexangulum Dentalium (LAMARCK) (LUKO­ Aequipecten macrotis ) badensis has a wide geogra­ . The stratigraphic (HÖRNES, 1856), from the Višnjica (PARTSCH in : : Fissidenta­ (KOJUM Fissiden Fissiden Ö RNES, Spi­ ­ ­ ­ ­ ­ ­

connection of the Central Paratethys with other marine areas. of connection Paratethys other with marine the Central had a wide geographic distribution, which implys an open marine research. questionable further age in and require is the Lower Badenian, while other Middle Miocene findings are and Herzegovina stored at the Natural History Museum Belgrade Paratethys. tral cene) deposits of the south of and southwestern margin the Cen Mio (Middle Badenian the from collected was material Fossil Museum. History Natural Croatian the and Belgrade Museum History Natural the at 1856) housed HÖRNES, (PARTSCHin evidence from the clayey sediments of Austria (Baden Austria of sediments clayey the from evidence fossilwithLower Badenian, the during widespreadspecieswas ied localities in Serbia, Bosnia and Herzegovina and Croatia. This mostly lived in muddy and muddy the south and southwestern margin of the Central Paratethys. constructionof the Middlefirst Miocenetransgression marine in detaileddentaliidreanalysis, the contribute to canfurther with NIĆ et al., 1995; JOVANOVIĆ, 2014).foraminifers(KOCHANSKY, These 1944; known STEVANOVIĆ, data, together 1977, AVA­ TEROT),pteropods ( accompaniedpelagicbya faunaofnautilids ( ljevik(Bogutovo Selo), togetherdentaliids,scaphopodswith are ment on the Medvednica Mt.), the wider Belgrade region and Ug Badenian)marlsofwidertheZagreb region („Čučerje“ develop al.,2012; LowerBadenian age(STUDENCKA etal., 1998; SELMECZI et of al., 1976; JOVANOVIĆ, 2014).austriaca These findings, as well as the record cochlear grade region, deep sea molluscs are very common: sidentalium mutabile SACCO, Zagreb) for the manuscript. help translating in Museum, History Natural (Croatian RADOVČIĆ Davorka Dr. to and paper, the improve to us helped which comments useful for reviewers anonymous and Zagreb) Museum, History tural versity of Zagreb, Croatia), Dr. Davor VRSALJKO (Croatian Na Herzegovina), Prof. Jasenka SREMAC (Faculty of Science, Uni Geology and Civil UniversityEngineering, of Tuzla, Bosnia and The Prof.authors thank Sejfudin VRABAC (Faculty of Mining, ACKNOWLEDGMENT badense This paper presents the scaphopod species 6. CONCLUSIONS RÖGL, 1998; STUDENCKA et al., 1998; KOVÁČ et al., 2007). („Transtethyan corridor“) andpassages marine the northern (e.g. of afore and corded re the to routes.Due migration and connections marine porary ROV, 1960). The distribution of (BALDI, 1960) and Bulgaria (KOJUMDGIEVA & STRACHIMI Slovakia (BALLUK, 1972; HARZHAUSER et al.,2007), Sus)(TITA, de 2011), (Lăpugiu 2009),Romania al., et (RÖGL Hungary F. badense avmsim duodecimlamellatum Parvamussium All the published data and records suggest that that suggest records and data published the All Lowerthe BadenianargillaceousIn marlsofwider Bel the According to the available fossil record, record, fossil available the to According species the Badenian, Lower the of part upper the During Serbia Bosnia and from age The of specimens analyzed the lived on a muddy to muddy POLI, JOVANOVIĆ, 2014). Miocene(probablythe Lower In KITTL and Dentalium to the area of Paratethys could be the southwestern Lentipecten corneus denudatus - mentioned findings, possible migration routes possible migration findings, mentioned ( Vaginellaaustriaca Dentalium (DODERLEIN) Aturia aturi F. badense ) michelottii michelottii -sandy substrate at all the stud (BASTEROT) (PET - sandy sandy substrate. . KITTL) and planktonic BON, niae a indicate (BRONN), Fissidentalium ba indicates the contem Geologia Croatica 69/2 Croatica Geologia (REUSS), HÖRNES and HÖRNES Aturiaaturi Fissidentalium Fissidentalium Neopycnodonte F.badense ­KOVIĆ et Vaginella - (BAS Sooss) ­dense Fis ­ ­ ­ ­ ­ ­ ­ ­ ­ ­ ­ ­ ­ ­

Geologia Croatica

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