Timing of Langhian bioevents in the Carpathian Foredeep and northern Pannonian Basin in relation to oceanographic, tectonic and climatic processes

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Geo log i cal Quar terly, 2018, 62 (1): 3–17 DOI: http://dx.doi.org/10.7306/gq.1399

Tim ing of Langhian bioevents in the Carpathian Foredeep and north ern Pannonian Ba sin in re la tion to ocean o graphic, tec tonic and cli ma tic pro cesses

Katarína HOLCOVÁ^1,*, Nela DOLÁKOVÁ², Slavomír NEHYBA² and František VACEK³

1 Charles Uni ver sity, In sti tute of Ge ol ogy and Palae on tol ogy, Albertov 6, 128 43 Praha 2, Czech Re pub lic

2 Masaryk Uni ver sity, In sti tute of Geo log i cal Sci ences, Fac ulty of Sci ence, Kotláøská 2, 611 37 Brno, Czech Re pub lic

3 Na tional Mu seum, Václavské námìstí 68, 115 79 Praha, Czech Re pub lic

Holcová, K., Doláková, N., Nehyba, S., Vacek, F., 2018. Tim ing of Langhian bioevents in the Carpathian Foredeep and north ern Pannonian Ba sin in re la tion to ocean o graphic, tec tonic and cli ma tic pro cesses. Geo log i cal Quar terly, 62 (1): 3–17, doi: 10.7306/gq.1399

The suc ces sion of bioevents in plank tonic foraminifer and cal car e ous nannoplankton com mu ni ties is re viewed and sum ma - rized for the Carpathian Foredeep and north ern Pannonian Ba sin in the time in ter val be tween ~16 and 13.5 Ma. This suc ces - sion can be sub di vided into three prin ci pal in ter vals: (1) an in ter val with rare Praeorbulina sicana and P. glomerosa. It was char ac ter ized by a lim ited im mi gra tion of in dex taxa linked to the lack of a warm sur face wa ter layer in the Cen tral Paratethys.

This in ter val can be cor re lated with the first Badenian trans gres sion near the Burdigalian/Langhian bound ary. The rare oc - cur rence of biostratigraphical mark ers does not al low its pre cise dat ing and in ter re gional cor re la tion; (2) a brief in ter val of the first oc cur rences of Praeorbulina circularis, Orbulina suturalis and Helicosphaera waltrans. This can be re lated to the for ma - tion of a warm sur face wa ter layer suit able for the sur vival of orbulinas and praeorbulinas and a change from estuarine to anti-estuarine cir cu la tion. This in ter val can be cor re lated with the sec ond Badenian trans gres sion, which, how ever, was not isochronous over the area as in ferred from dif fer ent suc ces sions of these first oc cur rences; (3) a lim ited ap pear ance of new in dex taxa in the Cen tral Paratethys prior to the Wielician Sa lin ity Cri sis. This time in ter val was char ac ter ized by in creased sea son al ity and sa lin ity os cil la tions fol lowed by cli mate cool ing. A “re verse” mi gra tion of the stress-tol er ant spe cies Helicosphaera walbersdorfensis from the Cen tral Paratethys to the Med i ter ra nean is sug gested. Sev eral lo cal bioevents with lim ited strati graphic cor re la tion po ten tial have been rec og nized in this in ter val.

Key words: Mio cene, Cen tral Paratethys, biostratigraphy, cal car e ous nannoplankton, foraminifera, palynology.

INTRODUCTION

The cor re la tion of the Langhian Stage with the lo cal Cen tral Paratethys stra tig ra phy has been re cently widely dis cussed.

Piller et al. (2007) cor re lated its base with the Karpatian/Badenian bound ary. This bound ary, de fined by the first oc cur rence of Praeorbulina (Papp et al., 1978), was sub se - quently re de fined by Hohenegger et al. (2014) to a level cor re - spond ing to the up per most Burdigalian. This cor re la tion was based on dat ing of the first oc cur rence of Praeorbulina in the world oceans. The top of the Langhian is cor re lated with the Moravian/Wielician lo cal substage bound ary (i.e., Early/Mid dle Badenian sensu Papp et al., 1978). Hohenegger et al. (2014) pro posed cor re la tion of the Moravian substage with the Early and Mid dle Badenian. How ever, this new sub di vi sion is not gen - er ally ac cepted and causes con fu sion in ter mi nol ogy and strati - graphic cor re la tions.

The Badenian ma rine biostratigraphy based on lo cal (eco)zones of Grill (1943) and Cicha et al. (1975) has been gen - er ally ac cepted since the 1970s (Papp et al., 1978). How ever, com ple men tary strati graphic data such as magnetostratigraphy (Hohenegger et al., 2009a; Selmeczi et al., 2012; de Leeuw et al., 2013) and new ra dio met ric and Sr-ages (Radócz, 2004;

Han dler et al., 2006; de Leeuw et al., 2010; Fordinál et al., 2014) showed that the con cept of such a seem ingly dis tinct sys - tem no lon ger ap plies even for the lo cal stra tig ra phy (Hohenegger et al., 2014).

De spite emerg ing new strati graphi cal meth ods and data, biostratigraphy re mains the most widely used cor re la tion method. The oc cur rence of com mon in dex taxa that mi grated in the sys tem of in ter con nected bas ins en able in ter re gional cor re - la tions of the Lower Badenian de pos its in the Cen tral Paratethys (Rögl, 1998; Popov et al., 2004). How ever, the tim - ing of such bioevents was not en tirely syn chro nous in the in di - vid ual bas ins with many fac tors, such as lo cal tec ton ics and cli - mate, wa ter chem is try and cir cu la tion re gime, play ing a role in their dis tri bu tion.

In this pa per we dis cuss the tim ing of the Langhian Cen tral Paratethys bioevents based on a case study of the north ern Pannonian Ba sin and the Moravian part of the Carpathian Foredeep. Our re sults are com pared to the world oceans

* Corresponding author, e-mail: holcova@natur.cuni.cz

Received: September 12, 2017; accepted: November 29, 2017; first published online: January 26, 2018

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and/or the Med i ter ra nean re gion and the re corded dif fer ences are re lated to cli ma tic, ocean o graphic and tec tonic fac tors that may have af fected the spa tial and tem po ral dis tri bu tion of in dex mi cro or gan isms.

GEOLOGICAL SETTING

Our study area in cludes the Pannonian Ba sin Sys tem (South Slo vak Ba sin) and the Carpathian Foredeep (Seneš, 1961; Fig. 1).

The South Slo vak Ba sin (SSB) be longs to the north east ern part of the Pannonian Ba sin Sys tem. The area was in un dated from the Oligocene to the Ottnangian (Mid dle Burdigalian) and then again dur ing the Karpatian (Late Burdigalian). Af ter the late Early Mio cene up heaval event, the sub si dence was re ju ve - nated around the Karpatian/Badenian (Burdigalian/Langhian) bound ary but only for a short time. Sed i men ta tion was con - trolled by NNW–SSE faults and was ac com pa nied by the Badenian ma rine trans gres sion. Depocentres were sit u ated in the west where the SSB con tin ued to the Dan ube Ba sin. Sub si - dence was fol lowed by a vol ca nic par ox ysm and ex ten sive vol - ca nism in ad ja cent ter res trial and ma rine ar eas. Vol ca nic bulg - ing caused rapid up lift and ma rine re gres sion in the area (Vass, 1995; Vass et al., 2007).

Sed i men ta tion in South Slovakia started with the tide-in flu - enced Príbelce Mem ber over lain by the Vinica For ma tion de - pos ited in lit to ral to neritic en vi ron ments. Tuffaceous sand - stones and siltstones with com mon bioturbation dom i nated by domichnia over lie the basal coarse volcaniclastic de pos its. Al - gal bioherms lo cally oc cur within the sand stones and siltstones (Vass et al., 2007).

The for ma tion and ba sin evo lu tion of the Carpathian Foredeep (CF) – as a pe riph eral fore land ba sin – was re lated to the subsurface load ing of the Al pine-Carpathian orogenic belt on the Bo he mian Mas sif mar gin dur ing the Early to Mid dle Mio - cene (Nehyba and Šikula, 2007; Nehyba et al., 2008). The ba - sin con tin ues south into the Al pine Molasse Zone and north into the Pol ish seg ment of the CF Ba sin (Oszczypko et al., 2006).

The infill and ba sin ar chi tec ture var ies through out the CF Ba sin, lo cal and re gional un con formi ties are de vel oped due to the vary ing in ten sity and ori en ta tion of flex ural load ing and dif fer ent geo log i cal and tec tonic his to ries of the base ment, along with a polyphase na ture of the ac tive ba sin mar gin and grad ual change of its po si tion (Brzobohatý and Cicha, 1993; Nehyba, 2000; Kováè et al., 2004; Oszczypko et al., 2006).

The strati graphic range of the sed i men tary infill of the stud - ied CF Ba sin seg ment is Eggerian to Lower Badenian (Brzobohatý and Cicha, 1993). The Lower Badenian de pos its re veal a dis tinc tive ba sin infill ge om e try be cause they are al - most sym met ri cally lo cated in the cen tral parts of the ba sin. The Lower Badenian sed i men tary se quence is dom i nated by a lithologically uni form pack age of pelitic sed i men tary strata (known as “Tegel”) with a thick ness reach ing sev eral hun dreds of metres. These pelites are in ter preted as de pos its of the mid - dle to outer shelf or even hemipelagites (Papp et al., 1978;

Cicha, 2001; Nehyba et al., 2008). Coarse-grained sand stones and con glom er ates rep re sent the sec ond dom i nant lithofacies be ing gen er ally de scribed as “basal or mar ginal coarse clastics”

(Krystek, 1974; Nehyba et al., 2008). Bioherms of red-al gal lime stones (Doláková et al., 2008) form lat er ally and vol u met ri - cally re stricted bod ies within the mudstones. Thin volcanoclastic beds (acidic tuffs and tuffites) in ter preted as dis - tal tephra fall out are rare (Nehyba et al., 1999).

MATERIALS AND METHODS

We syn the size multiproxy data from bore holes from the CF:

LOM-1 (Holcová et al., 2015a), ZIDL-1 and ZIDL-2 (Doláková et al., 2014), RY-1 (Kopecká, 2012), OV-1 and OV-2 (Nehyba et al., 2016); from the SSB (bore holes N-45, N-48, N-68, N-80, N-83, N-95, N-91; Holcová et al., 1996; for lo ca tion of all bore - holes see Fig. 1). In ad di tion, palynological data from the fol low - ing bore holes from the CF were used: IK-1 (Basistová and Doláková, 2011; Doláková et al., 2011), HJ-1, HJ-2, HJ-103 (Hladilová et al., 1999, 2001).

Foraminifera were ana lysed in the frac tion be tween 0.063 and 2 mm, cal car e ous nannoplankton was stud ied from smear slides. As sem blage quan ti ta tive eval u a tion was based on 200–300 spec i mens for foraminifera and 300–500 spec i mens for cal car e ous nannoplankton. The method of Zágoršek et al.

(2007) was used for our pur poses.

Stan dard mac er a tion in HCl (20%), HF, KOH and HCl (10%) and ZnCl2 (den sity = 2 g/cm³) was used for palynological sam ples. Pol len di a grams were pro cessed us ing POLPAL soft - ware (Walanus and Nalepka, 1999) with a min i mum of 150 de - ter mined pol len grains and spores, ex clud ing Pinus and un de - ter mined co ni fers. The ter mi nol ogy of Stuchlik et al. (1994), Kvaèek et al. (2006) and Kovar-Eder et al. (2008) was em - ployed for the clas si fi ca tion of veg e ta tion units.

We have used pub lished d¹⁸O data from foraminiferal tests for fur ther palaeoenvironmental in ter pre ta tions (Holcová and Demeny, 2012; Doláková et al., 2014; Scheiner, 2015). Four dif fer ent datasets have been dis tin guished , each char ac ter iz - ing spe cific lev els in the wa ter col umn: (1) data from Globigerinoides bulloides char ac ter ize sur face wa ters dur ing pe ri ods of en hanced pro duc tiv ity (prob a bly spring bloom;

Schiebel et al., 1997); (2) data from Globigerinoides char ac ter - ize sur face sum mer strat i fied wa ters (Reynolds and Thunell, 1985; Hemleben et al., 1989), (3) data from epifaunal Cibicidoides spp. re flect the qual ity of the bot tom wa ter (Kaiho, 1994; Murray, 2006); (4) pore wa ter chem is try in the sed i ment was doc u mented by iso to pic val ues from shal low infaunal Melonis spp. and Gyroidina spp. and deep infaunal Uvigerina spp. (Caralp, 1989; Hermelin, 1992; Sjoerdsma and Van der Zwaan, 1992; Sen Gupta and Machain-Castillo, 1993; Miao and Thunell, 1993; Rathburn and Corliss, 1994).

PALYNOLOGICAL ANALYSIS

Two prin ci pal zonal for est as sem blages have been rec og - nized from the study of palynoflora: (1) sub trop i cal broad-leaved for ests char ac ter ized by a high abun dance of ev - er green el e ments (rel a tive abun dance up to 38%), such as Sapotaceae, palms, Engelhardia, Platycarya, ev er green Fagaceae, Reevesia, Cornus-Mastixia, and Rutaceae; (2) a warm to tem per ate mixed mesophytic and broad-leaved de cid - u ous for est type with broad-leaved el e ments (rel a tive abun - dance 12–21%), i.e., Quercus, Celtis, Juglans, Tilia, Betula, and Acer. In creased di ver sity and quan tity of “de cid u ous oak type” pol len grains have been re corded here by con trast with the Lower Mio cene. Oc cur rences of moun tain for est com po - nents (Cedrus, Tsuga and Picea; Doláková et al., 1999, 2011, 2014; Kováèová et al., 2011), coastal swamp (Taxodiaceae, Cyrillaceae, Myricaceae, Decodon) and ri par ian el e ments (Alnus, Salix, Ulmus, Fraxinus, Liquid ambar, Carya) in di cate the com plex na ture of zonal biotopes in the ad ja cent ar eas.

The high est pro por tion of thermophilous flo ral el e - ments/low est per cent age of arctotertiary el e ments were de -

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Timing of Langhian bioevents in the Carpathian Foredeep and northern Pannonian Basin... 5

Fig. 1A – schematic map of the areas under study and their positions within the Carpatho-Pannonian region with location of the sections studied (map modified from Kováè et al., 2007);

B – chrono-, bio- and magnetostratigraphy of the interval studied

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tected in palynospectra from the in ter val with Praeorbulina sicana and P. glomerosa. A rel a tively lower abun dance of xerophilous el e ments such as Engelhardia and Olea has been ob served here (Fig. 2A).

An in crease in abun dance of arctotertiary el e ments that co - in cides with in creased abun dances of xerophilous mark ers (such as dry herbs and heliophytes Olea, Poaceae, Asteraceae, Caryophyllaceae, Chenopodiaceae, Ericaceae) can be seen in the short in ter val be tween the FOs of Praeorbulina circularis and Orbulina (Fig. 2B, C). The dis ap - pear ance of Sapotaceae and de crease in Cornoideae–Mas - tixioideae and ev er green Fagaceae and in crease in arctotertiary el e ments were re corded around the LCO of Helicosphaera waltrans.

The in ter val with Orbulina suturalis and com mon H.

walbersdorfensis is char ac ter ized by a dom i nance of co ni fers and ma rine dinoflagellates (some times re ferred to as the Pinus event; Fig. 2D). Pol len and spores com pletely dis ap pear higher in the sec tion in many of the bore holes stud ied.

BIOEVENTS SUCCESSION AND RELIABILITY OF THEIR USE IN STRATIGRAPHY

Sev eral prom i nent bioevents have been rec og nized in the bore holes and sec tions stud ied. Their po ten tial for lo cal and in - ter re gional biostratigraphy is dis cussed be low.

The first oc cur rences of Praeorbulina sicana and P.

glomerosa and the last oc cur rence of Helicosphaera ampliaperta. The in ter val be low the first oc cur rences (=FOs) of Helicosphaera waltrans, Praeorbulina circularis and Orbulina suturalis is char ac ter ized by scat tered oc cur rences of Praeorbulina sicana and P. glomerosa (Ap pen dix 1A*) that can - not be used for any re li able strati graphic cor re la tions.

Helicosphaera ampliaperta rarely oc curred in this in ter val; how - ever, the po si tion of its last oc cur rence (=LO) is in ac cu rate due to its scar city along the sec tions (Ap pen dix 1D). Its pos si ble redeposition must also be taken into con sid er ation.

The first oc cur rences of Praeorbulina circularis and Orbulina suturalis. The FOs of Praeorbulina circularis and Orbulina suturalis rep re sent a very dis tinct bioevent. Be sides the ap pear ance of new in dex taxa, the abun dance of plank ton from the Orbulina–Praeorbulina group is con sid er ably higher by con trast with the pre vi ous in ter val (Ap pen dix 1A).

The FO of Praeorbulina circularis slightly pre ceded the FO of Orbulina suturalis. Be fore this event, the Coccolithus pelagicus/Reticulofenestra minuta ra tio changed (Ap pen - dix 1B).

The last com mon oc cur rence of Helicosphaera waltrans and the last oc cur rence of Sphenolithus heteromorphus. Quan ti ta tive anal y ses of cal car e ous nannoplankton as sem blages showed in creased rel a tive abun - dance of H. walbersdorfensis (Ap pen dix 1C). The last com mon oc cur rence (=LCO) of H. waltrans co in cides with in creased abun dances of H. walbersdorfensis and can be used as a re li - able strati graphi cal marker (Ap pen dix 1C). This event oc curred af ter the FO of Orbulina. Above the LCO, H. waltrans has been re corded only dis con tin u ously in sev eral sam ples.

The LO of S. heteromorphus is not a re li able marker in the sec tions stud ied be cause of its scarce oc cur rences (Ap pen - dix 1D). Marunteanu (1999) and Bartol (2009) rec om mended us ing the de creased abun dance of Cyclicargolithus floridanus as an aux il iary in di ca tor in the Cen tral Paratethys. How ever,

this has not been rec og nized in our study ma te rial (Ap pen - dix 1E) nor have other syn chro nous bioevents de scribed (the LOs of discoasters, the FCO of Reticulofenestra pseudoumbilica >7 µm; Bartol, 2009).

Lo cal bioevents: acme of Reticulofenestra minuta, Spiroplectinella carinata and Globorotalia transylvanica.

The high abun dance of Reticulofenestra minuta is char ac ter is - tic of the stud ied sec tions above the FO of Helicosphaera waltrans (Ap pen dix 1B). An in crease in R. minuta abun dance co in cides with in creas ing num bers of H. walbersdorfensis (r = 0.74; p <0.001) and de crease in C. pelagicus. A neg a tive cor re - la tion of rel a tive abun dances of H. walbersdorfensis and C.

pelagicus spe cies can also be de tected (r = –0.56; p <0.001).

The lev els above the LCO of H. waltrans are char ac ter ized by higher vari a tions in R. minuta fre quency com pared to the un der - ly ing in ter val (Ap pen dix 1B).

An in crease in abun dance of biserial ag glu ti nated foraminifera (pri mar ily Spiroplectinella carinata) may be used for def i ni tion of a lo cal (eco)zone Spiroplectinella carinata (Grill, 1943) that cor re sponds to the Mid dle Badenian sensu Papp et al. (1978). The strati graphic dis tri bu tion of this morphogroup, sum ma rized in Ap pen dix 1F, showed that the in crease in abun - dance oc curred above the LCO of Helicosphaera waltrans.

How ever, this event has been rec og nized only lo cally (LOM-1, RY-1 and ZIDL-2 bore holes).

A G. transylvanica acme was re corded in the Pol ish and Ro - ma nian part of the CF prior to the Wieliczka sa lin ity cri sis. The gen er ally pos i tive trend in rel a tive abun dance of these en demic taxa with sev eral cy clic os cil la tions de tected in our sam ples (Ap pen dix 1G) may in di cate that a sin gle G. transylvanica acme may be only a lo cally re stricted event.

INTERREGIONAL CORRELATIONS AND BIOEVENTS TIMING

The first oc cur rences of Praeorbulina sicana and P.

glomerosa. In ac cor dance with our ob ser va tions, Praeorbulina sicana and P. glomerosa have been re corded only dis con tin u - ously through out the Cen tral Paratethys in the time in ter val be - fore the FO of Orbulina. Both spe cies have been rarely found in the Styrian Ba sin (Spezzaferri et al., 2009; Hohenegger et al., 2009a), the Al pine Foredeep Ba sin (Aus trian Mollase Ba sin;

Æoriæ et al., 2004), the Dan ube Ba sin (Rybár et al., 2015, 2016), the South Slo vak Ba sin (Vass et al., 2007) and the Pol ish part of the CF (Oszczypko and Oszczypko-Clowes, 2012). This in ter - val cor re sponds to lo cal biozone of Globigerinoides sicanus (Cicha et al., 1975).

Bioevent nu mer i cal dat ing in this lin eage for the Cen tral Paratethys is not gen er ally ac cepted due to the ab sence of re li - able ra dio met ric dat ing con trol. The tim ing of the praeorbulinas’

first oc cur rences in the Med i ter ra nean re gion mark edly dif fers from those in the world oceans (Ab dul Azis et al., 2008; Turco et al., 2011; Wade et al., 2011; Gradstein et al., 2012; Fig. 3). The use of bioevents for cor re la tion be tween the global magnetostratigraphical chart and de tected chrons in our study area may be mis lead ing. A pre sumed path way be tween the At - lan tic/Indo-Pa cific and the Cen tral Paratethys in this pe riod passed through the Med i ter ra nean (Rögl, 1999; Popov et al., 2004; Kováè et al., 2007, 2017a, b) to pres ent-day Slovenia and Croatia (this in ter val was re ported from this re gion by Cicha et al., 1975; Bartol, 2009). A strongly hy poth e sized di rect con nec - tion be tween the Paratethys and the Indo-Pa cific realm may

* Supplementary data associated with this article can be found, in the online version, at doi: 10.7306/gq.1399

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Fig. 2. Relative abundances of palaeoclimatological markers (A–D) terrestrial palynomorphs (E, F) d¹⁸O values (data from Holcová and Demeny, 2012;

Doláková et al., 2014; Holcová et al., 2015b; Scheiner, 2015)

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have led via the Cen tral Paratethys and the Transylvanian and Pannonian bas ins fur ther eastwards to the south ern mar gin of the Black Sea plate and the Pontids (Rögl, 1998; Fig. 4). How - ever, pos si ble ev i dence for this ma rine path way was de stroyed dur ing subduction of oce anic crust in the Al pine-Hi ma la yan orogenic belt.

For the rea sons de scribed above, we pre fer use of the Med - i ter ra nean bioevent dat ing for cal i bra tion of magneto - stratigraphical chrons de tected by Hohenegger et al. (2009a).

The FO of P. glomerosa in the world ocean dated at 16.4–16.1 Ma (Wade et al., 2011) cor re sponds to C5Cn.1n chron (16.29–15.97 Ma; Gradstein et al., 2012). If the nor mal po lar ity chron with the FO of P. glomerosa in the Styrian Ba sin (Hohenegger et al., 2009a) is cor re lated with the Med i ter ra nean da tum of the FO of the same taxon (15.2–15.1 Ma; di Stefano et al., 2008, Iaccarino et al., 2011), chron C5Bn.2n (15.15–15.04 Ma; Gradstein et al., 2012) may be in ferred (Figs. 3 and 4). Sim i larly, the re verse po lar ity chron with the FO of P. sicana can be in ter preted am big u ously based on a heterochronous tim ing of the FO of P. sicana in the world oceans (16.97 Ma based on Gradstein et al., 2012 or 16.4 Ma from Wade et al., 2011) and in the Med i ter ra nean (16.177 Ma;

Iaccarino et al., 2011; Turco et al., 2011). If the Med i ter ra nean age is ac cepted, the spe cies would ap pear in the Cen tral Paratethys in the ear li est Langhian C5Br Chron

(15.974–15.16 Ma) rather than in the lat est Burdigalian Chron C5Cn.1r (16.3 Ma; Gradstein et al., 2012; Figs. 3 and 4). More - over, praeorbulinas oc curred only dis con tin u ously in the sec - tions stud ied, in which case, de ter mi na tion of their ex act FOs may be mis lead ing and their oc cur rence can be used only for ap prox i mate dat ing of strata with P. sicana at ~15.9–15.1 Ma and with P. glomerosa from 15.1 to 14.6 Ma.

De pos its with rare Praerbulina sicana and P. glomerosa ap - pear to be re stricted to the Cen tral Paratethys. How ever, it is not ex actly known to what ex tent they rep re sent the orig i nal ma rine trans gres sion. The oc cur rence of re de pos ited mudstone intraclasts with the Langhian microfossils in de pos its of the sec - ond Badenian cy cle in di cates that it may have been re duced by sub se quent ero sion. On the other hand, they may have not been rec og nized in some ar eas due to a lack of rare praeorbulinas. Ad di tional nu mer i cal dat ing would cer tainly help to re solve these ques tions.

The first oc cur rences of Helicosphaera waltrans, Praeorbulina circularis and Orbulina suturalis. The FOs of Praeorbulina circularis, Orbulina suturalis and Helicosphaera waltrans rep re sent im por tant bioevents rec og nized over the Cen tral Paratethys. How ever, their suc ces sion may spa tially and tem po rally vary or be ab sent in in di vid ual Cen tral Paratethys bas ins (Fig. 5): O. suturalis with H. waltrans fre - quently ap pear si mul ta neously in the CF in Ukraine and in the Fig. 3. Correlation of magnetostratigraphical chrons detected in the Central Paratethys (Hohenegger et al., 2009a) with a global magnetostratigraphical chart based on the FOs of index taxa in the world oceans and the Mediterranean

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North Cro atian Ba sin, while praeorbulinas do not oc cur here (Æoriæ et al., 2009; Gozhyk et al., 2015). A H. waltrans oc cur - rence with out orbulinas has been re ported from the East ern Paratethys (Gozhyk et al., 2015). Orbulinas with out H. waltrans were de scribed from the north ern part of the Dan ube Ba sin (Rybár et al., 2015).

The FO of Orbulina suturalis of ten co in cides with the FOs of other praeorbulinas, pri mar ily P. circularis. How ever, in the Styrian Ba sin, the Al pine Foredeep Ba sin and the CF, the FO of P. circularis slightly pre ceded the FO of O. suturalis (Cicha et

al., 1975; Æoriæ et al., 2004; Spezzaferri et al., 2009; Doláková et al., 2014; Fig. 5).

Orbulinas are the most widely dis trib uted in dex microfossils in this time in ter val, which oc cur in all Cen tral Paratethys bas ins (e.g., Æoriæ et al., 2004, 2009; Tomanová-Petrová and Švábenická, 2007; Spezzaferri et al., 2009; Selmeczi et al., 2012; de Leeuw et al., 2013; Peryt, 2013; Doláková et al., 2014;

Holcová et al., 2015a, b; Gozhyk et al., 2015; Rybár et al., 2015). The FO of Orbulina rep re sents the base of the lo cal biozone of Praeorbulina-Orbulina suturalis (Cicha et al., 1975).

Fig. 4. Magnetostratigraphically calibrated FOs of Orbulina in the Central Paratethys (based on data of Æoriæ et al., 2004; Hohenegger et al., 2009a; de Leeuw, 2011; Selmeczi et al., 2012) with possible immigration

pathways (palaeogeographic reconstruction modified from Rögl, 1998 and Kováè et al., 2017a)

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The magnetostratigraphical tim ing of the O. suturalis FO dif - fers in in di vid ual bas ins (Fig. 4). In the Transylvanian Ba sin it is cal i brated at 14.6–14.17 Ma, which cor re sponds to the C5ADn Chron (de Leeuw, 2011). This is con sis tent with the Med i ter ra - nean dat ing (14.59 Ma; di Stefano et al., 2008; Figs. 3 and 4). In other bas ins, world-ocean bioevent ages for cal i bra tion of de - tected magnetostratigraphical chrons were used. In the Pannonian Ba sin, it ap peared in the re verse chron cor re lated with C5Bn.1r Chron (15.03–14.87 Ma; Selmeczi et al., 2012). A sim i lar age is sug gested for the Al pine Foredeep Ba sin (Æoriæ et al., 2004), while the spe cies’ first oc cur rence in the Styrian Ba - sin is dated to the nor mal Zone C5Bn.1n (14.87–14.78 Ma). In the case of Med i ter ra nean bioevents dat ing, de tected magnetochrons can be re-cal i brated to the C5ADn Chron as in the Transylvania Ba sin. This cal i bra tion can be also sup ported by ra dio met ric dat ing of ho ri zons close to the FO of Orbulina in Retznei Quarry: 14.21 and 14.39 Ma (Han dler et al., 2006). Re - verse chrons in the Al pine Foredeep and Pannonian bas ins can be cor re lated with C5ACr Chron (14.163–14.033 Ma) and may in di cate a grad ual ap pear ance of Orbulina in the Cen tral Paratethys from the south to the north.

The first oc cur rences of Helicosphaera waltrans and the last oc cur rences of Helicosphaera ampliaperta. Gen er - ally, last oc cur rences have lim ited cor re la tion po ten tial in epicontinental seas due to the ef fect of spe cific palaeo - environments, which may lo cally ac cel er ate or de lay spe cies ex tinc tion. Re work ing of microfossils may com monly oc cur here. This may ex plain vari able rel a tive po si tions of the LO of Helicosphaera ampliaperta in com par i son to other events (Fig. 5). In the world oceans and the Cen tral Paratethys this event oc curred af ter the FO of P. glomerosa. By con trast, a re - verse bioevent suc ces sion can be traced in the Med i ter ra nean.

The Med i ter ra nean spe cies Helicosphaera waltrans ap peared in the Cen tral Paratethys above the FO of P. glomerosa, si mul ta - neously with the LO of H. ampliaperta (Æoriæ et al., 2004; Bartol, 2009) or slightly later (Spezzaferri et al., 2009). The co-oc cur rence of both helicosphaeras (Švábenická, 2002) may re flect ei ther spe - cific con di tions in the CF en abling co-oc cur rence of both spe cies or redeposition of H. ampliaperta from older strata. The LOs of H.

ampliaperta may be cor re lated with the LOs of other large helicosphaeras (H. scissura, H. mediterranea; Švábenická, 2002) which sup ports the first pos si bil ity.

How ever, both events have lim ited ap pli ca bil ity in biostratigraphy: the FO of H. waltrans due to its scar city and the LO of H. ampliaperta due to its pos si ble re work ing.

Last com mon oc cur rence of Helicosphaera waltrans.

The tim ing of this eas ily rec og niz able event can be ap prox i - mated from dat ing of the Sooss bore hole (Vi enna Ba sin) at 14.38–14.14 Ma (Hohenegger et al., 2009b). H. waltrans was not re corded in this bore hole. In the Styrian Ba sin H. waltrans was re corded in a ra dio met ri cally dated ho ri zon (14.39 Ma;

Hohenegger et al., 2009a). This dat ing in di cates that the event oc curred at ~14.39–14.38 Ma, in ac cor dance with the Med i ter - ra nean dat ing of this event (14.357 Ma, Ab dul-Azis et al., 2008;

14.414 Ma, Hüsing et al., 2010).

The last oc cur rence of Sphenolithus heteromor phus.

The LO of S. heteromorphus is a sig nif i cant event in the Med i - ter ra nean (Hilgen et al., 2009) which is, how ever, poorly rec og - nized in the Cen tral Paratethys. Its strati graphic range may be re duced in some bas ins (e.g., the north ern part of the Dan ube Ba sin; Rybár et al., 2015).

Lo cal bioevents. Lo cal bioevents (acme of Reticulofenestra minuta, Spiroplectinella carinata, Globorotalia transylvanicaand miliolids) can only be traced in the Cen tral Paratethys. These events prob a bly re flected spe cific lo cal palaeoenvironments.

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CLIMATOSTRATIGRAPHIC AND SEQUENCE STRATIGRAPHIC CORRELATION

Climatostratigraphy. The very low abun dance of arctotertiary mark ers and high num ber of thermophylous el e - ments in the in ter val with the first praeorbulinas can be cor re - lated with the peak of the Mid dle Mio cene Cli ma tic Op ti mum (MMCO; 15 Ma; Gradstein et al., 2012). In creased abun dances of xe ro phyt ic mark ers in the same in ter val cor re spond with the first Med i ter ra nean aridification event dated at 15.074 Ma (Hüsing et al., 2010). These ages fit well to the cal i bra tion of magnetostratigraphical chrons us ing the Med i ter ra nean bio - event dat ing.

Sub se quent in crease in abun dance of arctotertiary mark ers in di cates cli mate cool ing around the FO of Orbulina; in creased hu mid ity can be in ferred from a de crease in xe ro phyt ic mark ers.

This change is fol lowed by grad ual cool ing (in creased abun - dance of arctotertiary mark ers) and aridification (in creased xe - ro phyt ic mark ers), which cor re sponds to the Mi-3a cool ing event (14.3 Ma; Gradstein et al., 2012) and more dis tinc tive Mi-3b event (13.8 Ma; Gradstein et al., 2012).

Se quence stra tig ra phy. Two depositional cy cles have been iden ti fied within the Moravian se quence of the CF. The first cy cle, cor re lated with TB2.3 of Haq et al. (1987), dates to the pre-Mid dle Badenian (Hohenegger et al., 2014). The sec - ond one, widely rec og niz able in the Paratethys Prov ince (TB2.4; Haq et al., 1987), cor re sponds to the Mid dle Badenian (Hohenegger et al., 2014). De pos its of the first depositional cy - cle have been pen e trated in bore holes in the Al pine-Carpathian Foredeep (Æoriæ and Rögl, 2004). Al though they have not been rec og nized in the out crops and shal low bore holes we stud ied, their re gional oc cur rence can be sup ported by the com mon pres ence of mudstone intraclasts (Eggenburgian to Early Badenian in age; Nehyba et al., 2006) re de pos ited in “the coarse-grained basal Badenian clastics” (Nehyba et al., 2008).

They are in ter preted as coarse-grained deltaic sys tem de pos its (Nehyba et al., 2008).

The sec ond Badenian depositional cy cle is mostly rep re - sented by outer shelf de pos its or hemipelagites. These de pos - its vol u met ri cally pre dom i nate in the part of the CF stud ied.

They can also be found fur ther west wards as scat tered ero - sional relicts doc u ment ing the orig i nal ex tent of these de pos its (Hladilová et al., 1999; Holcová et al., 2015a; Nehyba et al., 2016).

EARLY BADENIAN PALAEOENVIRONMENT AND ITS INFLUENCE ON BIOEVENT TIMING

Sev eral key fac tors in flu enced the dis tri bu tion of in dex taxa in the Cen tral Paratethys: (1) the qual ity of the Cen tral Paratethys wa ter-masses (i.e., tem per a ture, sa lin ity, nu tri ent and ox y gen con tent) played a role in the suc cess ful sur vival and re pro duc tion of in dex taxa; (2) the cir cu la tion re gime pri mar ily in the Med i ter ra nean–Paratethys sys tem (estuarine vs anti- estuarine, see Fig. 6). The anti-estuarine re gime may have trig - gered plank ton im mi gra tion, while pre dom i nantly estuarine cir - cu la tion may have had the op po site ef fect; (3) the ex is tence, char ac ter (mainly depth) and wa ter qual ity of com mu ni ca tion cor ri dors. Gen er ally, the greater the depths of the com mu ni ca - tion path way, the lesser the ef fect of cli mate, pres ence of other cor ri dors or cur rent re gime on plank ton ex change (this prin ci - pally ap plies for depths >1,000 m). In shal lower cor ri dors (<200 m), ex change with the ad ja cent oceans may be highly sen si tive to the above-men tioned fac tors (Vara, 2015). Since

the lat ter ar range ment is ex pected in our study area, the pre vi - ously de scribed fac tors must be taken into con sid er ation.

For in ter pre ta tion of cir cu la tion re gime, sea-wa ter strat i fi ca - tion may be sig nif i cant. The strat i fi ca tion was re con structed from sta ble ox y gen iso tope data for the in ter val above the FO of Orbulina. The foraminifera tests from the in ter val be low the FO of Orbulina were poorly pre served. The in ter val be tween the FO of Orbulina and the LCO of Helicosphaera waltrans is char ac - ter ized by sig nif i cant dif fer ences in sur face and bot tom wa ter ox y gen iso tope val ues (Fig. 2E, F), which may in di cate the pres ence of a well-strat i fied wa ter col umn. Sea sonal vari a tions in iso tope val ues be tween spring and sum mer can also be seen (Fig. 2E, F). This vari a tion is less pro nounced above the LCO of Helicosphaera waltrans, and can be in ter preted as a re sult of mix ing of sur face and bot tom wa ters dur ing the spring plank ton bloom and in creased sa lin ity/tem per a ture vari abil ity in sum mer.

In the fol low ing chap ter we dis cuss mi gra tion path ways and re stric tions of new or gan isms in the three strati graphic in ter vals dis tin guished.

In ter val with Praeorbulina sicana and P. glomerosa dom i nated by estuarine cir cu la tion (15.9–14.3 Ma). The cor ri - dor be tween the At lan tic and the Med i ter ra nean re mained rel a - tively deep dur ing the Early Langhian, which en abled en try of cold At lan tic wa ter as shown by the oc cur rence of psychrospheric ostracods (Benson, 1978). The palaeo bio logi cal ev i dence can be sup ported by car bon iso tope data (Vergnaud- Grazzini, 1983, 1985), which dem on strates iden ti cal iso tope com po si tion in the At lan tic and west ern Med i ter ra nean bot tom wa ters. Cen tral Med i ter ra nean estuarine cir cu la tion is sug gested for the Early Langhian up to the FO of Orbulina (up per part of the P. sicana Subzone, i.e. mid dle part of the MNN5a Subzone;

Dall’Antonia et al., 2001). The Indo-Pa cific con nec tion re mained open with the Circumequatorial Cur rent in place (von der Heydt and Dijkstra, 2005) in the time in ter val prior to the FO of Orbulina (Gebhardt, 1999). The pres ence of di a toms, high-nu tri ent Globigerina and Coccolithus pelagicus and high-nu tri ent ben thic foraminiferal taxa may also in di cate an estuarine cir cu la tion in the Cen tral Paratethys (Tomanová-Petrová and Švábenická, 2007).

The rare oc cur rence of Praeorbulina sicana and P.

glomerosa in the Cen tral Paratethys may be re lated to pre vail - ing estuarine cir cu la tion and re stricted plank ton im mi gra tion (Fig. 6). An other fac tor that lim ited Praeorbulina oc cur rence in the Cen tral Paratethys may be the lack of a suit able palaeo - environment for this plank ton group. Both Orbulina, and Praeorbulina thrived in strat i fied oceans with a sum mer warm wa ter layer in place (Chap man and Da vis, 2010). This can be ev i denced by the iden ti cal iso to pic com po si tion of both groups in our study ma te rial (Scheiner, 2015). This warm sur face wa ter layer was prob a bly ab sent in the Cen tral Paratethys dur ing the Early Badenian, which can also be doc u mented by the rare oc - cur rence of other warm-wa ter el e ments as Globigerinoides or Globigerinella. Helicosphaera ampliaperta and H. waltrans co-oc cur rence in the Cen tral Paratethys in con trast to the Med i - ter ra nean also in di cates a dif fer ent qual ity of sur face wa ters in both re gions. The ear lier dis ap pear ance of H. ampliaperta in the Med i ter ra nean at ~15.5 Ma (Iaccarino et al., 2011) may have ac cel er ated the evo lu tion of Helicosphaera waltrans in a Med i ter ra nean va cant niche.

The MMCO at ~15 Ma can be cor re lated with this in ter val (Zachos et al., 2001; Gradstein et al., 2012). How ever, stud ies on microflora and macroflora from the Pannonian and Vi enna bas ins showed some dif fer ences to the global trends over the Early and Mid dle Mio cene. The Early Langhian tem per a ture peak does not rep re sent the Mio cene tem per a ture max i mum in

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the study area. The warm est cli mate was re corded in the Eggenburgian–Ottnangian tran si tion (~17–16 Ma; Knobloch et al., 1975; Planderová, 1990; Jiménez-Moreno et al., 2005, 2006; Kvaèek et al., 2006; Doláková et al., 2011a, b; Kováèová et al., 2011; Doláková et al., 2014). This cor re sponds to the con clu sions of Böhme et al. (2007), who de scribed the suc ces - sion from paratropical ev er green for est be tween 17.5–17.3 Ma, fol lowed by sub trop i cal semi-de cid u ous and oak-lau rel sub trop - i cal for est (17–15.3 Ma) from the North Al pine Foredeep xyloflora. This prob a bly re flects mesoclimatic changes caused by north erly plate tec tonic shift and/or up lift of the Carpathian moun tain chain.

Tran si tion from estuarine to anti-estuarine cir cu la tion in the in ter val with Praeorbulina circularis, Orbulina suturalis and Helicosphaera waltrans (14.6–14.3 Ma). Clo - sure of the In dian Gate way in the Mid dle Langhian caused a tran si tion from estuarine to anti-estuarine cir cu la tion in the proto-Med i ter ra nean (Kouwenhoven, 2000). This can be dated to the time in ter val be tween the FO of O. suturalis and the FO of

O. universa (Russo et al., 2007) at 14.56–14.36 Ma (di Stefano et al., 2008). Change in the Med i ter ra nean cir cu la tion re gime also in flu enced cir cu la tion changes along the Med i ter ra - nean–Cen tral Paratethys com mu ni ca tion gate way and trig - gered sev eral dis tinct bioevents (i.e., FOs of Praeorbulina circularis, Orbulina suturalis and Helicosphaera waltrans over a short pe riod of time which co in cides with the drop in the Coccolithus pelagicus/Reticulofenestra minuta ra tio;

Tomanová-Petrová and Švábenická, 2007; Spezzaferri at al., 2009). This is in ter preted as a re sult of vari a tions in sur face wa - ter qual ity. C. pelagicus is an in di ca tor of cold and nu tri ent-rich wa ters (Okada and McIntyre, 1979; Win ter et al., 1994; Cachao and Moita, 2000), which is con sis tent with the estuarine cir cu la - tions in the Early Langhian. On the other hand, the oc cur rence of R. minuta in di cates en vi ron men tal stress char ac ter ized by sa lin ity and nu tri ent os cil la tions (Flores et al., 1997; Wells and Okada, 1997; Kameo, 2002; Wade and Brown, 2006). De te ri o - rat ing qual ity of sur face wa ters may be linked to sa lin ity os cil la - tions in a downwelling cir cu la tion re gime ex pand ing from the Fig. 6. Models of immigration of biostratigraphical index planktonic foraminifera

and calcareous nannoplankton to the Central Paratethys during the Early Badenian (Langhian)

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Med i ter ra nean to the Paratethys. Our de tailed anal y sis shows that the in ter val with H. waltrans rep re sents a tran si tion be - tween both re gimes with lo cally per sist ing coastal upwelling (Holcová et al., 2015a). Gen er ally, this newly es tab lished anti-estuarine cir cu la tion trig gered the for ma tion of a warm sum mer sur face wa ter layer and the im mi gra tion of plank tonic taxa such as orbulinas and praeorbulinas and also Globigerinoides spp. and Globigerinella regularis (Fig. 6;

Holcová et al., 2015a).

The var ie gated suc ces sion of the FOs of Praeorbulina circularis, Orbulina suturalis and Helicosphaera waltrans and the po si tion of the FO of Orbulina in nor mal and re verse magnetochrons sug gests that the sec ond Badenian trans gres - sion was not isochronous across dif fer ent bas ins. This ef fect is gen er ally rec og nized in pe riph eral fore land bas ins as the for ma - tion of ac com mo da tion space in dis tal and prox i mal parts (Heller et al., 1988; Catuneanu et al., 1997, 1998).

In ter val with Orbulina suturalis and com mon H.

walbersdorfensis dom i nated by anti-estuarine cir cu la tion and on set of the Wielician sa lin ity cri sis (14.3–13.8 Ma). No global bioevent has been rec og nized in the Cen tral Paratethys dur ing this in ter val. A lo cal event char ac ter ized by a grad ual sub sti tu tion of large H. waltrans by small H. walbersdorfensis oc curred at its base. The rel a tive abun dance of H.

walbersdorfensis pos i tively cor re lated with Reticulofenestra minuta (Holcová, 2017) sug gests its stress-tol er ance. The palaeoenvironmental changes dur ing this event were sum ma - rized by Doláková et al. (2014), Holcová et al. (2015a, b), and Nehyba et al. (2016), be ing char ac ter ized by in creas ing arid ity and de creas ing riverine and ter res trial nu tri ent in put with ep i - sodic heavy rain falls. In creased sea son al ity caused per tur ba - tions in mixed and strat i fied wa ter col umns and sea sonal vari a - tions in nu tri ent in put. In crease in the sur face wa ter sa lin ity dur - ing sum mers can be de tected in the ox y gen iso tope re cord (Scheiner, 2015) and dinoflagellate as sem blages (Nehyba et al., 2016). Aridification trends can also be traced in the in - creased per cent age of xe ro phyt ic mark ers in the pol len spec tra from our stud ied bore holes (Fig. 2C). A sim i lar trend dated at 14.3 Ma was de scribed by Böhme et al. (2010) from xyloflora of the Al pine Foredeep Ba sin.

The tim ing at ~14.3 Ma can also be ap prox i mated to the FCO of H. walbersdorfensis. The age of its FCO in the Med i ter - ra nean (14.05 Ma; Mourik et al., 2011) in di cates the im mi gra - tion di rec tion from the Paratethys to the Med i ter ra nean, which is ex pressed in the bioevents suc ces sion. The stress-tol er ant H.

walbersdorfensis, a typ i cal Paratethyan spe cies, re flected a vary ing qual ity of sur face wa ter in a small epicontinental ba sin which ex tended to the Med i ter ra nean due to the Mid dle Mio - cene Cli ma tic Tran si tion, while clo sure of the In dian–Med i ter ra - nean Gate way and a change in the cir cu la tion pat tern also oc - curred in the At lan tic–Med i ter ra nean Gate way at the end of the Langhian (Gebhardt, 1999).

This in ter val is char ac ter ized by a dom i nance of co ni fers in the pol len spec tra, which is some times re ferred to as the Pinus event. The ac cu mu la tion of co ni fer pol len in off shore ma rine sed i ments may be ex plained by pol len mass pro duc tion, long-dis tance ae rial trans port (from W or NW) and also by their high re sis tance to ox i da tion in wa ter or sed i ment (Heusser, 1978; Hopkins and Mc Car thy, 2002). A cy clic ar range ment of this mass ac cu mu la tion is con sis tent with vari a tions in abun - dance of biserial ag glu ti nated foraminifera and Globorotalia transylvanica/bykovae and also with multiproxy cyclicity in the Sooss bore hole (Vi enna Ba sin), in ter preted as Milankovitch cli - ma tic cy cles (Hohenegger et al., 2008).

The in ter val was ter mi nated by the Wielician sa lin ity event that led to in creased sa lin ity of the sur face wa ter layer (Scheiner, 2015; Nehyba et al., 2016). This can be doc u mented by an in crease in miliolid abun dance in the shal low-wa ter de - pos its. This bioevent is dated at 13.73 Ma in the Dan ube Ba sin (Fordinál et al., 2014) which is con sis tent with dat ing of the sa - lin ity cri sis (be gin ning at 13.81 Ma with a du ra tion be tween 200 and 600 ky; de Leeuw et al., 2010).

Al though the persistant anti-estuarine cir cu la tion should gen - er ally fa vour the im mi gra tion of biostratigraphic plank ton mark ers to the Cen tral Paratethys (the FO of G. praemenardi at 14.4 Ma in the world oceans or 13.9 Ma in the Med i ter ra nean; Gradstein et al., 2012), the cool ing (re corded in mi cro- and macroflora) and sa lin ity os cil la tions in the sur face wa ters prob a bly re stricted this pro cess. The sa lin ity cri sis started prior to the LO of S.

heteromorphus in the world oceans (13.5 Ma; Gradstein et al., 2012), which sug gests that the bioevent oc curred ear lier in the Cen tral Paratethys due to os cil lat ing sa lin ity.

Sa lin ity os cil la tions ac com pa nied by salt de po si tion can be de tected all over the Med i ter ra nean re gion (evaporite de pos its in Egypt; Ied et al., 2011) and may be used as a cor re la tion ho ri - zon in se quences with no in dex taxa.

The cool ing event can be de tected slightly later in the con ti - nen tal plant as sem blages (at ~13.5 Ma; Planderová, 1990;

Jiménez-Moreno et al., 2005; Jiménez-Moreno, 2006; Kvaèek et al., 2006; Doláková et al., 2011, 2014; Kováèová et al., 2011). This dis pro por tion has not yet been ad e quately ex - plained.

CONCLUSIONS

1. The in ter val be tween the FO of Praeorbulina and the FO of H. waltrans and/or Orbulina cor re lates with the first Badenian trans gres sion. Prob a bly only the Med i ter ra nean–Cen tral Paratethys Gate way was in place at this time with no di rect con - nec tion be tween the Cen tral Paratethys and the world ocean.

There fore, the tim ing of biostratigraphical events re corded here must be co eval with or later than in the Med i ter ra nean. In this case, cor re la tion of the Karpatian/Badenian bound ary with the Burdigalian/Langhian bound ary (in the sense of Piller et al., 2007) is ap pro pri ate, and can be cor rob o rated by climatostrati - graphic data plac ing the top of the MMCO to this in ter val.

The first Badenian biostratigraphical mark ers (Praeorbulina sicana and P. glomerosa) were only rarely de tected in the Cen - tral Paratethys. This is in ter preted as a con se quence of re stricted im mi gra tion of the in dex taxa linked to the ab sence of a warm sur face wa ter layer in this re gion and pre dom i nantly estuarine cir cu la tion in the Med i ter ra nean–Paratethys sys tem. The scar - city or ab sence of stan dard biostratigraphical mark ers and/or pres ence of nu mer ous gaps re lated to tec tonic up lift com pli cate strati graphic cor re la tion in this in ter val, and more ra dio met ric ages would cer tainly help un der stand ing of this pe riod.

2. The co eval FOs of Praeorbulina circularis, Orbulina suturalis and Helicosphaera waltrans at the base of the sec ond Badenian transgressive cy cle co in cide with the change from estuarine to anti-estuarine cir cu la tion re gime in the Med i ter ra - nean–Paratethys sys tem. This was ac com pa nied by es tab lish - ment of a warm sum mer sur face wa ter layer in the Cen tral Paratethys. Slightly dif fer ent bioevent tim ings and their suc ces - sion may in di cate heterochrony of the trans gres sion in in di vid - ual bas ins and the for ma tion of a sum mer sur face wa ter layer.

3. The Middle Mio cene cli ma tic tran si tion sig nif i cantly af - fected the Cen tral Paratethys palaeoenvironment in the time in -

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ter val fol low ing the LCO of Helicosphaera waltrans. This re - sulted in cli ma tic in sta bil ity, and cli mate cool ing fol lowed by a re gional sa lin ity cri sis. Its on set can be de tected ear lier in ma - rine set tings and later in con ti nen tal en vi ron ments. The ab - sence of stan dard strati graphi cal mark ers can be re lated to palaeoenvironmental fluc tu a tions and sa lin ity os cil la tions.

Ac knowl edge ments. The study was sup ported by the pro - jects PROGRES Q45, Min is try of Cul ture programme DKRVO 2016/04 (Na tional Mu seum, 00023272). The au thors greatly ap pre ci ate the con struc tive re views of D. Peryt and M. Oszczypko-Clowes which im proved the manu script.

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