New biostratigraphic ev i dence (cal car e ous nannofossils, ostracods, foraminifers, ammonites, inoceramids) on the Mid dle Coniacian
in the east ern Bo he mian Cre ta ceous Ba sin
Andrea SVOBODOVÁ
1, 2,*, Tomáš KOÈÍ
3, Martina KOÈOVÁ VESELSKÁ
1, 2, Bruno FERRÉ
4, Mar tin KOŠ¸ÁK
2, Stanislav ÈECH
5and Markéta CHROUSTOVÁ
2, 51 In sti tute of Ge ol ogy of the Czech Acad emy of Sci ences, Rozvojová 269, 165 00, Prague 6, Czech Re pub lic
2 Charles Uni ver sity in Prague, Fac ulty of Sci ence, Albertov 6, 128 43, Prague 2, Czech Re pub lic
3 Na tional Mu seum, Nat u ral His tory Mu seum, Palaeontological De part ment, Cirkusová 1740, Prague 9 – Horní Poèernice, Czech Re pub lic
4 Dame du Lac 213, 3 rue Henri Barbusse, F-76300 Sotteville-lÀs-Rouen, France
5 Czech Geo log i cal Sur vey, Klárov 3/131, 118 21, Prague 1, Czech Re pub lic
Svobodová, A., Koèí, T., Koèová Veselská, M., Ferré, B., Košïák, M., Èech, S., Chroustová, M., 2019. New biostratigraphic ev i dence (cal car e ous nannofossils, ostracods, foraminifers, ammonites, inoceramids) on the Mid dle Coniacian in the east - ern Bo he mian Cre ta ceous Ba sin. Geo log i cal Quar terly, 63 (3): 586–602, doi: 10.7306/gq.1489
As so ci ate Ed i tor – Micha³ Zatoñ
The clas si cal lo cal ity of Svinary in the east ern Bo he mian Cre ta ceous Ba sin is the site of new biostratigraphic in ves ti ga tions.
Be sides some scarce macrofossil ev i dence, bulk sed i ment sam ples were pro cessed to re trieve micropalaeontological as - sem blages, and cal car e ous nannofossil smear slides were ana lysed. The stud ied ma te rial pro vided cal car e ous nannofossil as sem blages in clud ing Micula staurophora, Lithastrinus septenarius and Broinsonia parca expansa, thus doc u ment ing the Mid dle Coniacian (up per part of UC10 Zone and lower part of UC11 Zone). The foraminifera as sem blage is rel a tively rich, plank tonic spe cies show a wide strati graphi cal range, while the ben thic as so ci a tion rep re sented by Neoflabellina suturalis suturalis and Gaudryina carinata is very sim i lar to the Coniacian biozone of Stensioeina granulata-Eponides whitei, valid for the Bo he mian Cre ta ceous Ba sin. Ostracods are rep re sented by two com mon cytherellid spe cies, and two rare or na mented spe cies: Imhotepia marssoni? and Pterygocythereis spinosa. Inoceramid bi valves, namely Platyceramus mantelli, and a newly re corded ammonite, Tridenticeras tridens, sup port the late Mid dle Coniacian age of the Svinary out crop. New biostratigraphic re sults are given along with palaeo eco logi cal in ter pre ta tions of newly col lected fos sil ma te rial.
Key words: biostratigraphy, Coniacian, Late Cre ta ceous, Bøezno For ma tion, Bo he mian Cre ta ceous Ba sin, Czech Re pub lic
INTRODUCTION
The Up per Cre ta ceous strata of the Bo he mian Cre ta ceous Ba sin (BCB) have been in tensely stud ied by nu mer ous au - thors (i.e. Krutský et al., 1975; Èech et al., 1980; Èech, 1989, 2011; Èech and Švábe nická, 1992, 2017; Ulièný et al., 1997, 2009, and oth ers); the stra tig ra phy based on fau nal as sem - blages of these se quences is the core of sev eral pa pers (Èech and Švábenická, 1992, 2017; Košïák et al., 2004; Svobodová
et al., 2014; Nádaskay et al., 2017). Al though they pro vided im por tant and rel e vant in for ma tion on the Cre ta ceous Sys tem of the BCB, the Mid dle Coniacian de pos its have been less thor oughly stud ied than older strata (e.g., Èech and Švábenická, 1992; Lees, 2008). In 2016 and 2017, field works were con ducted on the Svinary out crop where the Mid dle Coniacian suc ces sion is best ex posed. The out crop pro vided a con tin u ous bi otic re cord al low ing a bed-by-bed col lect ing of nu mer ous stratigraphically sig nif i cant fos sils, such as cal car e - ous nannoplankton, fora minifers, inoceramid bi valves, and rare ammonites. Hereby, we in tro duce a new re cord of a tridenticerid heteromorph ammonite within the BCB. This pa - per pres ents new data to re con struct the palaeoenvironment of these fau nal as sem blages, and sub se quently to get a more pre cise age of the Svinary de pos its. The new fos sil re cord def - i nitely con trib utes to a better knowl edge of the Mid dle Conia - cian suc ces sion in the east ern part of the BCB (Fig. 1).
* Corresponding author, e-mail: asvobodova@gli.cas.cz Received: December 13, 2018; accepted: July 20, 2019; first published online: October 2, 2019
GEOLOGICAL AND GEOGRAPHICAL SETTINGS
The clas si cal lo cal ity of Svinárky was first men tioned by Frič (1889). This lo cal ity is sit u ated on the right bank of the Orlice River in the vi cin ity of Svinary, close to the house No. 19, east of Hradec Králové (Fig. 2A). The nat u ral out crop it self, show ing soft grey cal car e ous claystones (Fig. 2B), ex - ceeds 15 m in its over all thick ness (Fig. 3). Frič (1889) as - signed the lo cal ity to the Up per Turonian of the Teplice lay ers.
Later, Zahálka (1949) dated this sec tion as the Up per Turonian (zones Xa, b, c) Scaphites Zone, men tion ing the fol - low ing spe cies: Mesocrinus fischeri (Geinitz), Ostrea vesicu - laris (Lamarck), Leda semilu naris (von Buch), Inoceramus cuvieri Sowerby (non Goldfuss), and Inoceramus costellatus Woods. Soukup (1962) was the first who at trib uted the Svinary de pos its to the Coniacian up per se ries of Xea with ar gil la - ceous pelosiderite con cre tions. He chal lenged Zahálka’s (1949) iden ti fi ca tion of the inoceramid fauna, es pe cially that of Inoceramus costellatus Woods.
Soukup (1962) men tioned in his guide (p. 90): “... From the wider area around Svinárky are known out crops of the for ma - tion with pelosiderite con cre tions (Xea), which be longs to the Inoceramus koeneni Zone; in a num ber of sur round ing out - crops are also oc cur ring small, ir reg u larly spher i cal and cy lin dri - cal clay-phos phate con cre tions (with about 15% P2O5) in car - bon ate claystones, which are very char ac ter is tic of the same fa - cies of the Březno lay ers in var i ous re gions of the Bo he mian Cre ta ceous Ba sin...”.
So far, the biostratigraphy of this lo cal ity has been based on macrofossils, es pe cially inoceramid bi valves that were stud ied in de tail for both tax o nomic and strati graphic pur poses. Nev er - the less, other biostratigraphic el e ments (mi cro- and nanno - fossils) have been put aside. There fore, the pres ent study aims to com pile new fos sil ma te rial from the Svinary out crop to up - date the biostratigraphy of the stud ied sec tion.
MATERIAL AND METHODS
CALCAREOUS NANNOFOSSILS
Cal car e ous nannofossils were ana lysed in smear slides pre pared by the decantation method us ing a 7% so lu tion of H2O2 (e.g., Švábenická, 2012) and mounted in Entellan. A to tal of 15 sam ples were ex am ined un der an Olym pus BX51 light mi - cro scope us ing an im mer sion ob jec tive with a 100× mag ni fi ca - tion. Dig i tal im ages of nannofossil spec i mens were made us ing an Olym pus DP70 dig i tal cam era. In or der to ob tain the rel a tive nannofossil abun dances and semi-quan ti ta tive in for ma tion about the re spec tive cal car e ous nannofossil as sem blages, 500 spec i mens were counted in each slide. Biostratigraphic data were then in ter preted with ref er ence to the UC zones of Bur nett (1998). Smear slides are now stored in the palaeontological col - lec tions of the Na tional Mu seum in Prague, un der reg is tra tion num ber NM-d27/2018.
OSTRACODS, FORAMINIFERS AND ASSOCIATED MACROFAUNA
For micropalaeontological pur poses, bulk rock sam ples were col lected ev ery 1 m from the base of the pro file up to its top on the right side of the nat u ral out crop. Sam ples were also taken from sev eral lev els: ~200 g sam ple for foraminifers, an other one for ostracods. Bulk sam ples had been chem i cally pro cessed with a 10% ace tic acid so lu tion for 24 hours and with a 10% hydrogene per ox ide for 1 hour, and then washed and sieved through a 0.063 mm mesh-sized sieve. Sub se quently, fos sils from this frac tion were sorted, hand-picked and iden ti fied un der a Bresser bin oc u lar mi cro scope (20× to 40× mag ni fi ca tion).
Pho tos of microfossil ma te rial were taken us ing a scan ning elec tron mi cro scope (SEM Hitachi S-3700N) in low vac uum.
Macrofossils were pho to graphed un der an gled light us ing a
Fig. 1. Palaeogeographic po si tion of the Bo he mian Cre ta ceous Ba sin (Up per Cre ta ceous) and the Svinary lo cal ity (marked with ar row)
Mod i fied af ter Košïák et al. (2018)
Canon EOS 550D dig i tal cam era. In view of the clayey char ac - ter of the sed i ment, coat ing of spec i mens with am mo nium chlo - ride for pho tog ra phy to en hance con trast deemed un fea si ble.
Spec i mens were mea sured us ing the microphotography set ting of the Olym pus DP70. Mea sure ments are given in milli metres (mm). All mi cro- and macrofaunal ma te rial is de pos ited in the palaeontological col lec tions of the Na tional Mu seum in Prague, un der reg is tra tion num bers NM-O8455-62.
RESULTS
CALCAREOUS NANNOFOSSILS
The sam ples pro vide mostly mod er ately pre served and rel - a tively abun dant nannofossil as sem blages. In to tal, 68 cal car e - ous nannofossil taxa were iden ti fied. De tailed in for ma tion about Fig. 2. Geo graph ical lo ca tion of the Mid dle Coniacian lo cal ity of Svinary within
the Bo he mian Cre ta ceous Ba sin and Eu rope (A); pic ture of the Svinary sec tion (B)
nannofossil dis tri bu tion and abun dance is given in Ta ble 1. The most sig nif i cant com po nents of the as sem blage (20–30%) is Watznaueria barnesiae (Fig. 4S), fol lowed by Micula stauro - phora and Broinsonia parca expansa. Other quan ti ta tively sig - nif i cant taxa are Eiffellithus turriseiffellii, Zeugrhabdotus scu - tula, Z. diplogrammus, and Helicolithus trabeculatus. Less fre - quently, but con tin u ously oc cur ring spe cies are Marthasterites furcatus, Micula concava, Gartnerago obliquum, Kamptnerius
magnificus, Prediscosphaera cretaceae, Lucianorhabdus male formis, Eiffellithus eximius and Lithraphidites carniolensis.
Rare and ir reg u lar oc cur rences of stratigraphically sig nif i cant taxa (e.g., Lithastrinus septenarius, L. grillii, Lucianorhabdus arcuatus and Zeugrhabdotus biperforatus) were also ob served (Figs. 4 and 5). A list of cal car e ous nannofossil taxa in al pha bet - i cal or der is given in Ap pen dix 1*.
Fig. 3. Strati graphic and lith o logic pro file of the Mid dle Coniacian lo cal ity of Svinary
* Supplementary data associated with this article can be found, in the online version, at doi: 10.7306/gq.1489
OSTRACODS
The suc ces sive ostracod as sem blages in this Mid dle Conia cian lo cal ity are gen er ally poor in both spec i mens and spe cies. Ev ery ostracod sam ple con tains a max i mum of four spe cies, namely Cytherella gr. ovata (Roemer, 1841), Cythe - rella cf. parallela (Reuss, 1846), Imhotepia marssoni? (Po - korný, 1984) and Pterygocythereis spinosa (Reuss, 1846) (see Fig. 6). Pokorný (1979) pro vided a sche matic ta ble of ostracod ranges for the Bo he mian Cre ta ceous Ba sin and erected eight ostracod as sem blage zones. His youn gest ostracod zone Cd cor re sponds to the lower part of the Inoceramus (Volvicera - mus) involutus Zone.
The ge nus Imphotepia, with a spe cial con cern to Imhotepia marssoni?, ranges from the Up per Turonian to the Coniacian in the BCB (Damotte et al., 1981). The rel a tively poor state of pres - er va tion ham pers any fur ther subspecific iden ti fi ca tion (Fig. 6D).
FORAMINIFERS
Among all sam ples from the Svinary lo cal ity, 10 planktic and 24 ben thic foraminifera spe cies have been indentified. The foraminiferal as sem blage con sists of di verse ben thic and
plank to nic spec i mens with some smaller, prob a bly ju ve nile forms (Figs. 7 and 8A, B). Foraminifera shells are rel a tively poorly pre served, white to light grey, filled by micrite, in some cases by py rite. Shell walls are of ten frag men tary or dis solved;
some shells are flat tened as a re sult of de for ma tion caused prob a bly by diagenetic al ter ations.
The foraminiferal as sem blage in cludes ben thic cal car e ous [e.g., Globorotalites gr. micheliniana (d’Orbigny, 1840), Trita - xia sp., Praebulimina sp., Neoflabellina suturalis suturalis (Cus - h man, 1935), Frondicularia apiculata Reuss, 1844], ag glu ti - nated [Spiroplectammina praelonga (Reuss, 1845), Gau dry ina carinata Franke, 1914, see Ap pen dix 2], and plank tonic spe cies [Planoheterohelix globulosa (Ehrenberg, 1840), White inella brittonensis (Loeblich and Tappan, 1961), W. baltica Douglas and Rankin, 1969, W. archaeocretacea Pessagno 1967, Dicarinella canaliculata (Reuss, 1854), D. imbricata (Mor nod, 1950), Marginotruncana pseudolinneiana Pessagno, 1967, and Marginotruncana coronata (Bolli, 1945)].
The plank tonic foraminiferal as sem blage is dom i nated by forms with glob u lar cham bers, rep re sented by Whiteinella spe - cies. Among dou ble-keeled foraminifera pre vail gen era Margino truncana and Dicarinella, the only rep re sen ta tive of biserial plank tonic forms is Planoheterohelix globulosa (Ehren - berg, 1840).
Chronostratigraphy, nannofossil zonation and ver ti cal dis tri bu tion
A – abun dant (more than 100 spec i mens per sam ple); C – com mon (11–100 spec i mens per sam ple);
The ben thic foraminiferal as sem blage is more di verse than the plank tonic one and in cludes stratigraphicaly im por tant spe - cies of Gaudryina carinata Franke, 1914 and Neoflabellina suturalis suturalis (Cushman, 1935) for BCB (Hercogová, 1982;
Hradecká, 2012).
Be cause of poor pres er va tion of foraminifera shells (caused prob a bly by diagenetic al ter ations) it was not al ways pos si ble to de ter mine all shells to the spe cies rank; some spe cies are iden - ti fied to the ge neric level.
AMMONITES
Two frag ments of the stratigraphically im por tant hetero - morph ammmonite Tridenticeras tridens (Schlüter, 1876) (Ancylo ceratina; Nostoceratidae) have been col lected. The larger one (spec i men No. NM-O8455) is ~25 mm high and con - sists of eight (prorsiradiate) ribs, three rows of prom i nent tu ber - cles (lat eral to mar ginal; Fig. 9A) and one row of weakly de vel - oped um bil i cal tu ber cles (Fig. 9B, spec i men No. NM-O8456).
The sec ond and third rows of tu ber cles are ar ranged in sep a - rated pairs (Fig. 9A). Main bi fur cate ribs start at um bil i cal tu ber - cles (Fig. 9B) and con tinue to mar ginal tu ber cles. Non-tu ber cu - late sin gle ribs (1–2) in ter ca late the tu ber cu lar ones; they are slightly con cave and weakly de vel oped. The su tures are not
pre served. The FO of Tridenticeras tridens is doc u mented in the mid dle/up per part of the Gauthiericeras margae Zone, i.e.
the up per Mid dle Coniacian (see dis cus sion be low).
INOCERAMIDS
One spec i men of the spe cies Platyceramus mantelli (de Mercey, 1872) was found 3 m above the base of the Svinárky out crop. The spec i men has a slightly pointed beak, oval rugae and growth lines. This out line char ac ter ized the morphotype beyenburgi Seitz, 1965 (Fig. 9C). The sec ond bro ken in com - plete spec i men with a costellatus-like or na ment can not be de - ter mined pre cisely, but it re sem bles Inoceramus frechi Flegel, 1905 (Fig. 9D).
ASSOCIATED MACROFOSSILS
Infaunal bi valves are very scarce, only two spec i mens of de - posit-feed ing nuculids, Nuculana (Jupiteria) semilunaris (Buch) and Nymphalucina laminosa (Reuss, 1846), were found in the out crop, rep re sent ing the Nucula – Nuculana as sem blage (Čech and Švábenická, 1992). Infaunal ir reg u lar echinoid tests of Hemiaster (Leymeriaster) cf. regulusanus (d’Orbigny, 1854) are more fre quent but com monly squashed. This echinoid
T a b l e 1 of cal car e ous nannofossils in stud ied sam ples
F – few (3–10 spec i mens per sam ple); R – rare (1–2 spec i mens per sam ple); ? – ques tion able iden ti fi ca tion; cf. – con fer
taxon be came spe cial ized to muddy sub strates in rather deeper ma rine set tings, point ing to off shore deeper wa ters (Fraaije et al., 2018).
The oc cur rence of araucarid gym no sperm plant leaves of Pagiophyllum brachyphyllum (Bayer) Kunzmann, 2007, con - sti tutes an ex cep tional find and sup ports prox im ity to the land (Fig. 8I).
NANNOFOSSIL BIOSTRATIGRAPHY
The biostratigraphically sig nif i cant spe cies are as fol lows:
Lithastrinus septenarius, L. grillii, Zeugrhabdotus biperforatus, Marthasterites furcatus, Broinsonia parca expansa, Luciano - rhabdus arcuatus, Micula staurophora and M. concava. Rare oc - cur rences of Micula swastica, Quadrum gartneri and Octo lithus multiplus were also no ticed. Helicolithus turonicus is ab sent.
Micula staurophora com monly oc curs across the en tire pro - file. The first oc cur rence (FO) of this spe cies de fines the base of the UC10 Nannofossil Zone in the lower Mid dle Coniacian (Bur - nett, 1998; Švábenická, 2012). The fol low ing UC11 Zone is de - fined by the FO of Lithastrinus grillii. Ir reg u lar re cords of this stratigraphically sig nif i cant spe cies were ob served from sam ple No. 4 up the sec tion (see Ta ble 1). Based on the pres ence of Micula staurophora, Lithastrinus septenarius and Broinso - nia parca expansa, sam ples Nos. 1–3 be long to the UC10 Zone. Co-oc cur rence of these taxa along with Lithastrinus grillii from sam ple No. 4 doc u ments the UC11 Zone, more ac cu rately the UC11a–b Subzones sensu Bur nett (1998). The marker spe cies for the base of the UC11c Nannofossil Subzone in the Up per Coniacian, Lucianorhabdus cayeuxii, has not been found.
DISCUSSION
Rel a tively nu mer ous stud ies based on cal car e ous nanno - fossils fo cused on the Lower and Up per Coniacian, and spe cif i - cally on the Turonian-Coniacian and Coniacian -Santonian boun d a ries (e.g., Melinte and Lamolda, 2007; Kędzierski, 2008;
Lees, 2008; Blair and Watkins, 2009; Svobodová et al., 2014).
The pres ent study deals with Mid dle Coniacian ma te rial. As far as biostratigraphy is con cerned, the stud ied sam ples range from the UC10 to UC11 zones. Bur nett (1998) placed the base of the UC11 Zone, de fined by the FO of Lithastrinus grillii, in the Up per Coniacian. Hampton et al. (2007) shifted this bioevent into the Mid dle Coniacian in the chalk fa cies of south ern Eng - land. Such a po si tion within the Mid dle Coniacian has also been ac cepted by Švábenická and Bubík (2014) for the Bo he mian Cre ta ceous Ba sin and the Outer West ern Carpa thians. Ac cord - ing to Švábenická et al. (2016) and Čech and Švábenická (2017), sam ples yield ing Lithastrinus grillii, Luciano rhabdus arcuatus and Marthasterites furcatus are in ter preted as the up - per part of the Mid dle Coniacian. More over, the con tin u ous pres ence of Lithastrinus septenarius ex cludes any over lap into the UC12 Zone (see Ta ble 1).
The con tent of Marthasterites furcatus in the cal car e ous nannofossil as sem blage does not ex ceed 2% in each sam ple, which also ex cludes the Marthasterites furcatus acme de - scribed across the Turonian-Coniacian bound ary (e.g., Švábe - nická, 2009, 2010; Švábenická and Bubík, 2014; Švábenická and Havlíček, 2017).
Dur ing the de po si tion of the Mid dle Coniacian sed i ments of the Březno For ma tion (Xe: sensu Soukup, 1956), the most suit - able con di tions for a mixed (plank tonic and ben thic) fora mini -
feral as sem blage pre vailed. The oc cur rence of dou ble-keeled plank tonic foraminifers (Dicarinella and Marginotruncana gen - era) doc u ments a well-ox y gen ated in ter me di ate-to-deep wa ter col umn (Štemproková-Jírová, 1978), while sur face wa ters are in hab ited by glob u lar forms such as the ge nus Whiteinella (Keller et al., 2001). The mode of life strat e gies is nei ther fully un der stood yet, nor easy to sum ma rize.
None the less, Premoli Silva, Sliter (1999) and Petrizzo (2002) and other au thors ap ply the eco log i cal con cept of K- and r-strat e gists for Cre ta ceous plank tonic foraminifera.
While globotruncanids and marginotruncanids are de scri - bed as K-strat e gists, Archaeglobigerina cretacea (d’Orbigny, 1840) and the ge nus Whiteinella are more r-se lected in ter me di - ate strat e gists (Petrizzo, 2002), Planoheterohelix globulosa (Ehren berg, 1840) is con sid ered an op por tun ist and r-strat e gist (Petrizzo, 2002) with tol er ance to lower ox y gen lev els (Keller at al., 2001). How ever, Abramovich et al. (2003) in ter preted hetero helicids as in hab it ants of subsurface lay ers, or wa ter masses close to the thermocline. Fur ther more, Premoli Silva, Sliter (1999) men tioned that heterohelicids rep re sented the dom i nant el e ment in the plank tonic foraminiferal as sem blage of the open ocean wa ters in the Early Cre ta ceous.
Among plank tonic spe cies, marginotruncanids (Margino - truncana pseudolinneiana, Marginotruncana coronata), Archaeo globigerina cretacea and dicarinellids (Dicarinella cana liculata, D. imbricata and other spe cies – see above) are char ac ter is tic for this lo cal ity. The plank tonic foraminifera as - sem blage be longs to the Dicarinella concavata Biozone sensu Robaszynski and Caron (1995) and cor re sponds with work of Štemproková-Jírová (1969, 1978), valid for the BCB.
The large ben thic spe cies of Neoflabellina suturalis sutu - ralis (Cushman, 1935) is typ i cal for the up per part of the Lower Coniacian, and ranges into the Up per Coniacian in the BCB (Hercogová, 1982; Hradecká, 2012). The oc cur rence of Neo - flabellina suturalis suturalis cor re sponds to its oc cur rence in the Lower Coniacian Buchletein Mem ber of the Sandbach For ma - tion of south east ern Ger many (Schnei der et al., 2011). These au thors com pared it with the stan dard chalk sec tion of Läger - dorf (Schönfeld, 1990: 135) in north ern Ger many, where Neofla bellina suturalis suturalis has its FO in the Mid dle Conia - cian. Like wise, the re spec tive FO of Neoflabellina suturalis ssp.
in the Münsterland Cre ta ceous Ba sin, north west ern Ger many (Dölling et al., 2014: 528; Fig. 4), lies within the up per part of the UC10 nannofossil zone. None the less, Niebuhr et al. (1999) re - port the FO of Neoflabellina (in clud ing the Neoflabellina sutu - ralis group) from the Mid dle Coniacian in the ex tra-Al pine Cre ta - ceous lay ers of Ba varia (Ohmert, 1969) and Aus tria (Wessely et al., 1981).
The ben thic foraminifera as sem blage shows sim i lar ity to the fauna struc ture of the Stensioeina granulata-Eponides whitei ben thic foraminifera biozone, based on sum mary re search from the Bo he mian Cre ta ceous Ba sin (Hradecká, 2012). For this biozone, a di verse ben thic as sem blage with abun dant ag glu ti - nated spe cies of the ge nus Gaudryina (e.g., G. carinata Franke, 1914) and cal car e ous ben thic spe cies, such as Neoflabellina suturalis suturalis (Cushman, 1935), is char ac ter is tic. The Stensio eina granulata-Eponides whitei Biozone is clas si fied as Lower to Up per Coniacian (Hradecká, 2012). Gyrodinoides nitidus (Reuss, 1844), wide spread in Up per Cre ta ceous epi - continental de pos its, seems not to be very use ful as a palaeo - environmental proxy; it is prob a bly not a very sen si tive spe cies to en vi ron men tal changes (Dubicka et al., 2014).
Pokorný (1979) and Damotte et al. (1981) did not men tion Cytherella gr. ovata, Cytherella cf. parallela and Pterygo cythe - reis spinosa. Fol low ing the re vi sion of the ge nus Pterygo - cythereis from the BCB by Pokorný (1966, 1987), this ge nus
Fig. 4. Cal car e ous nannofossils of the Mid dle Coniacian lo cal ity of Svinary
A – Ahmuellerella octoradiata, sam ple 4; B – Biscutum ellipticum, sam ple 10; C – Biscutum melaniae, sam ple 2; D – Eiffellithus eximius, sam ple 1; E – Eiffellithus gorkae, sam ple 4; F – Eiffellithus turriseiffelii, sam ple 5; G – Broinsonia enormis, sam ple 1; H – Broinsonia signata, sam ple 10; I – Broinsonia parca expansa, sam ple 15; J – Rhagodiscus angustus, sam ple 15; K – Cylindralithus biarcus, sam ple 6; L – Cribrosphaerella ehrenbergii, sam ple 12; M, N – Helicolithus trabeculatus, sam ples 14 and 1; O – Chiastozygus litterarius, sam ple 9; P – Retecapsa crenulata, sam ple 6; Q – Retecapsa octofenestrata, sam ple 8; R – Staurolithites ellipticus, sam ple 5; S – Watznaueria barnesiae, sam ple 14; T – Watznaueria bri tan nica, sam ple 5; U – Reinhardtites anthophorus, sam ple 13; V – Tetrapodorhabdus decorus, sam ple 3; X, Y – Manivitella pemmatoidea, sam ples 6 and 15; Z – Prediscosphaera cretacea, sam ple 10; AA – Prediscosphaera ponticula, sam ple 7; AB – Prediscosphaera columnata, sam ple 10; AC – Prediscosphaera spinosa, sam ple 9; AD – Prediscosphaera cf;
grandis (sensu Bur nett 1998), sam ple 4; AE – Tranolithus orionatus, sam ple 9; AF, AG – Marthasterites furcatus, sam ples 10 and 1; AH – Gartnerago obliquum, sam ple 13; AI, AJ – Kamptnerius magnificus, sam ples 4 and 15; AK – Grantarhabdus coronadventis, sam ple 2;
cross-po lar ized light; scale bars – 5 mm
Fig. 5. Cal car e ous nannofossils of the Mid dle Coniacian lo cal ity of Svinary
A, B – Zeugrhabdotus biperforatus, sam ples 7 and 9; C – Zeugrhabdotus diplogrammus, sam ple 6; D – Zeugrhabdotus bicrescenticus, sam ple 11; E – Zeugrhabdotus erec tus, sam ple 2; F – Zeugrhabdotus embergeri, sam ple 2; G – Zeugrhabdotus scutula, sam ple 11; H – Quadrum gartneri, sam ple 1; I, J – Micula staurophora, sam ples 3 and 6; K, L – Micula concava, sam ples 3 and 7; M – Micula cubiformis, sam ple 15; N – Micula cf. swastica, sam ple 15; O, P – Micula adumbrata, sam ples 4 and 11; Q, R – Lithastrinus septenarius, sam ples 5 and 13; S, T – Lithastrinus grillii, sam ples 4 and 9; U, V – Eprolithus moratus, sam ples 2 and 4; X, Y – Eprolithus floralis, sam ple 10 (Y – side view); Z – Octolithus multiplus, sam ple 6; AA – Calculites ovalis, sam ple 9; AB – Calculites obscurus, sam ple 2; AC – Acuturris scotus, sam ple 11; AD – Lucianorhabdus maleformis, sam ple 9; AE – Lucianorhabdus quadrifidus, sam ple 3; AF, AG – Lucianorhabdus arcuatus, sam ples 8 and 10; AH – Lithraphidites carniolensis, sam ple 10; AI – Braarudosphaera bigelowii bigelowii, sam ple 3; AJ – Thoracosphaera operculata, sam ple 9; AK – Zeugrhabdotus scutula, two spec i mens, sam ple 14; cross-po lar ized light; scale bars – 5 mm
ranged from the Mid dle Turonian to the Coniacian. Pterygo - cythereis spinosa (Reuss, 1846) is men tioned from the Up per Turonian de pos its of Úpohlavy (Houdková, 2016). Based on the sculp ture and shape of car a pace for palaeo eco logi cal com - par i son, the spec i mens at hand of the ge nus Pterygocythereis may well com pare with ex tant Pterygocythereis spe cies: e.g.
Pterygocythereis (Pterygocythereis) jonesii (Baird, 1850) and Pterygocythereis ceratoptera (Bosquet, 1852). P. jonesii oc - curs in the Med i ter ra nean be tween 20 to 300 m (Puri et al., 1965; Breman, 1976; Peypouquet and Nachite, 1984), and in
the Adri atic Sea be tween 40 and 100 m (Breman, 1976) and be tween 80 and 170 m (Bonaduce et al., 1975). Pterygo - cythereis jonesii is a com mon sublittoral ma rine spe cies found at all depths down to 200 m. It is es sen tially a warm-wa ter form liv ing in the Med i ter ra nean (e.g., Zakynthos: Tsourou et al., 2012; Aegean Sea and Black Sea: Perçin-Paçal et al., 2015;
ÖzuluÈ et al., 2018; Sea of Marmara: Tunoglu, 1996, ÖzuluÈ et al., 2018; and the At lan tic con ti nen tal shelf of Spain: Pascual et al., 2008). Its most septentrional re cord lies in the Kattegat and Skagerrak (Wilkinson, 2007). Pterygocythereis ceratoptera is Fig. 6. Ostracods from the Mid dle Coniacian lo cal ity of Svinary
A, C – Cytherella gr. ovata (Roemer, 1841), scale bar – 500 mm; B – Cytherella cf. parallela (Reuss, 1846), scale bar – 300 mm; squashed valves; D – Imhotepia marssoni? (Pokorný, 1984), scale bar – 300 mm;
E – Pterygocythereis spinosa (Reuss, 1846), scale bar – 300 mm
Fig. 8. Se lected microfossils and macrofossils from the Svinary lo cal ity
A – Ramulina aculeata d’Orbigny, 1840, scale bar – 400 mm; B – Dorothia filiformis (Berthelin), scale bar – 500 mm;
C – Calcisphere (Cal car e ous dinocysts) spe cies Calcisphaerula innominata Bonet, 1986, scale bar – 50 mm; D – bore hole in an ostracoda test, scale bar – 500 mm; E – Porifera gen. et sp. indet., scale bar – 500 mm; F – frag ment of ophiuroid arm, scale bar – 100 mm; G – bourgueticrinid brachial plate, scale bar – 500 mm; H – bourgueticrinid brachial plate, scale bar – 500 mm; I – Araucarid gym no sperm plant leaves Pagiophyllum brachyphyllum (Bayer) Kunzmann, 2007, scale bar – 1 cm
Fig. 7. Foraminiferal as sem blage from the Mid dle Coniacian lo cal ity of Svinary
Ben thic cal car e ous foraminifera: A – Lagena cf. apiculata (Reuss, 1850), scale bar – 100 mm; B – Dentalina gracilis (d’Orbigny, 1840), scale bar – 200 mm; C – Nodosaria oligostegia (Reuss, 1845), scale bar – 400 mm; E – Lenticulina lobata (Reuss, 1845), scale bar – 300 mm; G – Gyrodinoides globulosa (Hagenow, 1842), scale bar – 1 mm; J – Praebulimina sp., scale bar – 100 mm; L – Frondicularia apiculata Reuss, 1844; scale bar – 400 mm; N – ?Eggerellina sp., scale bar – 500 mm; O – Lagena hispida (Reuss, 1858), scale bar – 300 mm; Q – Praebulimina reussi (Mor row, 1931), scale bar – 200 mm; R – Neoflabellina suturalis suturalis (Cushman, 1935), scale bar – 400 mm. Ben thic ag glu ti nated foraminifera: D – Spiroplectammina praelonga (Reuss, 1845), scale bar – 400 mm; M – Gaudryina carinata Franke, 1914, scale bar – 500 mm.
Planktic foraminifera: F – Planoheterohelix globulosa (Ehrenberg, 1840), scale bar – 200 mm; H – Archaeglobigerina cretacea (d’Orbigny, 1840), scale bar – 400 mm; H1 – Archaeglobigerina cretacea (d’Orbigny, 1840), scale bar – 300 mm; I – Marginotruncana marginata (Reuss, 1845), scale bar – 400 mm; K – Dicarinella canaliculata (Reuss, 1854), scale bar – 500 mm; K1 – Dicarinella canaliculata (Reuss, 1854), lat eral view, scale bar – 300 mm; P – Whiteinella brittonensis (Loeblich and Tappan, 1961), scale bar – 400 mm
re ported from the Aegean Sea and the Sea of Marmara (Tunoglu, 1996; ÖzuluÈ et al., 2018). Its up per bathymetric limit var ies in dif fer ent re gions, os cil lat ing be tween 2 m in some la - goons to 115 m in the Gulf of Biscay (Pokorný, 1987). This is re - lated mainly to the ac tual depth of the wave base or, more pre - cisely, the storm wave base (Liebau, 1984). Pterygocythereis has not been found liv ing above this level. Liebau (1984) des ig -
nated a ptery gocline – up per level of the bathymetric range of waves-avoid ing or gan isms. This con clu sion by Lieabau (1984) is sup ported by the fact that Pterygocythereis spe cies are gen - er ally not found in strand-line ma te rial, and that worn spe cies do not oc cur in autochthonous ma te rial.
The or na mented spec i mens in this ostracod as sem blage are rather small and de void of any oc u lar tu ber cle, thus in di cat - Fig. 9. Se lected macrofossils from the Svinary lo cal ity
A – Tridenticeras tridens (Schlüter, 1876), com pressed part of turriliticone whorl (adult); B – Tridenticeras tridens (Schlüter, 1876), com pressed part of turriliticone whorl (ju ve nile, ad o les cent); C – Platyceramus mantelli (de Mercey, 1872), morphotype beyenburgi Seitz, 1965. Lat eral view of the right valve; D – Inoceramus sp. aff. Inoceramus frechi Flegel, 1904; E – "Tro chus“ sp.;
F – Nuculana (Jupiteria) semilunaris (Buch). Lat eral view of the left valve; G – Nucula sp.; H – Hemiaster (Leymeriaster) cf.
regulusanus (d’Obirgny, 1854); all scale bars – 1 cm, ex cept G – 0.5 cm
ing a lim ited ox y gen short age within the first centi metres of mud-sup ported bot tom sed i ments (infaunal hab its).
Palaeoecologic con di tions of coccolithophorids are typ i cally con strained by the photic zone. The con tin ued pres ence of Lucianorhabdus with in creas ing oc cur rences start ing at sam ple 8 in di cates con tin u ous shallowing in the up per part of the pro - file. This is in ac cor dance with the co-oc cur rences of the Braarudo sphaera and Thoracosphaera nannofossil gen era, both doc u ment ing a po ten tial in put of terrigenous ma te rial (e.g., Wyton and Bown, 2007; Švábenická, 2010). Si mul ta neously, joint oc cur rences of the co ni fer Frenelopsis alata and araucarid gym no sperm plant leaves of Pagiophyllum brachyphyllum in sam ple 13 sup port the prox im ity to the land as well.
The FO of the ammonite Tridenticeras tridens is re ported from the up per part of the Prionocycloceras iberiense Zone in Spain and is cor re lat able to the FO in Ger many (Westphalia:
Kaplan and Ken nedy, 1994; Küchler, 1998). Kaplan and Ken - nedy (1994) re ported its strati graphic range from the up per Mid - dle Coniacian (Gauthiericeras margae ammonite Zone, Volvicera mus koeneni and Volviceramus involutus inoceramid Zones) through the Paratexanites serratomarginatus Zone in Ger many. In Spain, the strati graphic range is re ported from the mid dle/up per part of the Gauthiericeras margae Zone, up per part of the Prionocycloceras iberiense Zone, in clud ing the Tridenticeras II Event (Küchler, 1998). Tridenticeras tridens oc - curs above the FO of Tridenticeras varians (Schlüter, 1876), and the LO is re ported from the Hemitissotia turzoi Zone (up per Paratexanites serratomarginatus Zone through the low er most Santonian: Santamaría-Zabala, 1992; Santamaría-Zabala and Ricardo, 1993; Küchler, 1998).
Vašíèek (1992) es tab lished a new tridenticerid taxon (Tridenti ceras soukupi Vašíèek, 1992) based on well-pre - served ma te rial from the Mid dle Coniacian of Štíty (east ern part of the BCB, Peroniceras tridorsatum = P. subtricarinatum and Gauthie riceras margae Zones). The new taxon is char ac - ter ized by the pres ence of four rows of tu ber cles (in con trast to the sim i lar spe cies of T. varians with three tu ber cle rows).
How ever, the closely re lated spe cies Tridenticeras tridens has also four rows of sim i larly de vel oped tu ber cles. Thus, the dif - fer ences be tween T. soukupi and Tridenticeras tridens are slight; both spe cies may be conspecific. In such a con text, Tridenticeras soukupi re quires re vi sion and needs to be com - pared in more de tail to Tridenticeras tridens. The ac com pa ny - ing ammonite fauna re ported by Vašíèek (1992) is al most iden ti cal to those of Tridenti ceras tridens from Spain and Ger - many (Santamaría-Zabala, 1992; Kaplan and Ken nedy, 1994;
Küchler, 1998 and ref er ences therein). The inoceramid spe - cies Platyceramus mantelli and Volviceramus koeneni co-oc - cur and both are com monly ac cepted as mark ers for the base of the Mid dle Coniacian (Wala szczyk and Wood, 2018).
Platyceramus mantelli ranges up into the Up per Coniacian.
Up to now, no study deals with the strati graphic value of the sub spe cies or morphotypes of Platy ceramus mantelli. In the ad ja cent bore holes (VY-1 Všestary and Tb-1 Tøebechovice – re cent re vi sion of SÈ), the FO of the gen era Volviceramus – Platyceramus ap pears 75–100 m be low the pres ent sur face, ap prox i mately 50–60 m above the top of the Rohatce Mem ber (low er most Coniacian). Thus, the sur face out crop in Svinary falls in the youn ger part of the Mid dle Coniacian.
CONCLUSIONS
The clas si cal lo cal ity of Svinary in the east ern Bo he mian Cre ta ceous Ba sin has been re in ves ti gated within the scope of
ac quir ing new palaeontological data, im prov ing its bio strati - graphic as sign ment, and pro vid ing firm palaeoenvironmental ev i dence.
Mod er ately pre served and abun dant cal car e ous nanno - fossil as sem blages pro vided ev i dence for the Mid dle Coniacian age, namely the up per part of the Mid dle Coniacian. From the co-occurences of Micula staurophora, Broinsonia parca expan - sa, Lithastrinus septenarius, and L. grillii, the up per part of the UC10 Zone and lower part of the UC11 Zone sensu Bur nett (1998) are con firmed.
Based on its orig i nal re cord in Svinary, the strati graphic oc - cur rence of ammonite Tridenticeras tridens is in ac cor dance with its oc cur rences in north ern Spain and Westphalia (i.e. the up per part of the Gauthiericeras margae Zone). The Mid dle Coniacian age of the Svinary out crop is also sup ported by the occurence of the inoceramid spe cies Platyceramus mantelli as a Euramerican Bo real el e ment.
The eco log i cal groups and morphotypes in plank tonic foraminifera re corded in the Svinary lo cal ity per mit spec u la tions about the depositional en vi ron ment, but the palaeo ec ol ogy of Up per Cre ta ceous foraminifera is not en tirely clear from avail - able lit er a ture. None the less, the dou ble-keeled plank tonic dica - rinellids and marginotruncanids in hab ited prob a bly a well- ox y - gen ated in ter me di ate-to-deep wa ter col umn, whereas the glob - u lar plank tonic whiteinellids in hab ited sur face wa ters in this lo - cal ity. The abun dance of dou ble-keeled dicarinellids and margino truncanids is sim i lar through the geo log i cal pro file (see Fig. 4). Heterohelicids in hab ited subsurface lay ers or wa ters close to the thermocline, with their in creas ing abun dance in the mid dle part of the pro file in the Svinary out crop (see Fig. 4).
Ostracods are rep re sented mostly by the ge nus Cytherella (C. gr. ovata, C. cf. parallela), as well as by Imhotepia marsso - ni?, Bairdia gr. cretacea and Pterygocythereis spinosa. Cal car - e ous dinocysts in clude Calcisphaerula innominata. The macro - fossil ma te rial con sists of ammonites (Tridenticeras tridens), bi - valves (Platyceramus mantelli), echinoids [Hemiaster (Leyme - riaster) cf. regulusanus], ophiuroids and trochiid gas tro pods, as well as by rep re sen ta tives of the Nucula – Nuculana bi valve level-bot tom as sem blage. Fos sil plant re mains were also found (Pagiophyllum brachyphillum, Frenelopsis alata). The inoce - ramid and ammonite fau nas clearly in di cate a Mid dle Coniacian age for the en tire ex po sure at Svinary and doc u ment the youn - gest pre served de pos its of the muddy fa cies of the cen tral part of the BCB.
The fau nal as sem blages and the sed i men tary fa cies in di - cate a rel a tively low-en ergy hy dro dy namic re gime be low the storm wave base (Pterygocythereis ostracod), soft pelitic ground (suit able for, e.g., ir reg u lar echinoids), warm tem per a - ture, and a wa ter depth not ex ceed ing 200 m. The foraminiferal and nannoplankton com mu ni ties also doc u ment ris ing tem per - a tures. Abun dant plant re mains re corded in the up per part of the sec tion may in di cate a shallowing pro cess and/or a higher ter res trial in put to the ba sin.
Ac knowl edge ments. The au thors wish to ex press their most sin cere grat i tude to K. Holcová (Charles Uni ver sity) for loan ing screens and her valu able ad vice on micropalaeonto - logical meth ods, to J.W.M. Jagt (Nat u ral His tory Mu seum, Maastricht) for echinoid iden ti fi ca tion, to Z. Vašíèek (Czech Acad emy of Sci ences, In sti tute of Geonics) for as sis tance in de - ter min ing ammonite spec i mens, and to J. Kvaèek (Na tional Mu - seum, Prague) for as sis tance in iden ti fy ing plant spec i mens. The tech ni cal ex per tise of L. Váchová and J. Sklenáø (Na tional Mu - seum, Prague) in SEM im ag ing is grate fully ac knowl edged. T.K.
thanks P. Rajlich and R. Hampl for their valu able help dur ing field work in June 2016. The au thors would like to thank re view ers
M.s Wilmsen and Z. Dubicka for their valu able com ments. This work is a con tri bu tion to the Re search plan of the In sti tute of Ge - ol ogy of the Czech Acad emy of Sci ences, v.v.i., No. RVO67985831. It was fi nan cially sup ported by the Cen ter for Geosphere Dy nam ics (UNCE/SCI/006), PROGRES Q45, the
Min is try of Cul ture of the Czech Re pub lic (IP DKRVO 2019/2.I, Na tional Mu seum, 00023272), Grant Agency ČR No. 17-10982 S, and Grant Agency of Charles Uni ver sity, pro ject No. 204217.
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