GeolCarp_Vol70_No2_113

GEOLOGICA CARPATHICA

, APRIL 2019, 70, 2, 113–134

doi: 10.2478/geoca-2019-0007

www.geologicacarpathica.com

Revisiting the age of Jurassic coral bioherms

in the Pieniny Klippen Belt (Western Carpathians)

on the basis of benthic foraminifers

DARIA K. IVANOVA



, JÁN SCHLÖGL

, ADAM TOMAŠOVÝCH

, BERNARD LATHUILIÈRE

and MARIÁN GOLEJ

Geological Institute, Bulgarian Academy of Sciences, Acad. G. Bonchev Str., Bl. 24, 1113 Sofia, Bulgaria;



dariaiv@geology.bas.bg

Department of Geology and Paleontology, Faculty of Natural Sciences, Comenius University, Mlynska dolina, Ilkovičova 6,

842 15 Bratislava, Slovakia; jan.schlogl@ uniba.sk

Earth Science Institute, Slovak Academy of Sciences, Dúbravská cesta 9, 840 05 Bratislava, Slovakia; geoltoma@savba.sk, geolmgol@savba.sk

Université de Lorraine CNRS, GeoRessources Lab. BP 70239, 54506 Vandoeuvre-lès-Nancy, France; bernard.lathuiliere@univ-lorraine.fr

(Manuscript received September 13, 2018; accepted in revised form March 12, 2019)

Abstract: Coral bioherms of the Vršatec Limestone that formed massive, several tens of meters thick complexes during

the Jurassic were important sources of carbonate production, with carbonate sediment exported to deeper parts of

the Pieniny Klippen Basin (Western Carpathians). However, the age of these carbonate factories remains controversial.

New analyses of benthic foraminiferal assemblages occurring in coral bioherms and peri-biohermal deposits of

the Vršatec Limestone at five sites in the western Pieniny Klippen Belt (Vršatec-Castle, Vršatec-Javorníky, Malé Hradište,

Malé Hradište-Kalvária, and Drieňová Hora) show that these sediments were deposited during the Bajocian and were

lateral equivalents of crinoidal limestones and breccias, in contrast to previous studies suggesting that they were deposited

during the Oxfordian. First, all sites are characterized by similar composition of foraminiferal assemblages on the basis

of presence–absence data, although foraminiferal assemblages in biosparitic facies at Vršatec are dominated by miliolids

whereas biomicritic facies at Malé Hradište are dominated by the spirillinid Paalzowella. The composition of fora mi-

niferal assemblages does not differ between the lower and upper parts of the Vršatec Limestone. Second, foraminifer

species that were assumed to appear for the first time in the Oxfordian already occur in the Middle Jurassic sediments

of the northern Tethyan shelf. Third, the first and last appearances of foraminifers documented in other Tethyan regions

are in accordance with stratigraphic analyses and ammonoid occurrences, demonstrating that bioherm-forming coral

communities developed on the Czorzstyn Ridge during the Bajocian. Several species of foraminifers of the Vršatec

Limestone appeared for the first time during the middle or late Aalenian (Labalina occulta, Paalzowella feifeli) and

during the Bajocian (Hungarillina lokutiense, Radiospirillina umbonata, Ophthalmidium caucasicum, O. terquemi,

O. obscurum, Paalzowella turbinella, Cornuspira tubicomprimata, Nubecularia reicheli) or appeared for the last time in

the Bajocian (Tethysiella pilleri) or Early Bathonian (Ophthalmidium caucasicum, O. obscurum). The composition and

diversity of communities with benthic foraminifers of the Vršatec Limestone is similar to the composition of foraminiferal

communities on carbonate platform environments with corals of the French Jura and Burgundy during the Bajocian.

Keywords: Middle Jurassic, Bajocian, benthic foraminifers, coral reefs, Vršatec Limestone, Pieniny Klippen Belt,

Western Carpathians, Slovakia.

Introduction

Coral bioherms and peri-biohermal deposits in the Carpathian

realm were largely restricted to the western sector of the Czor-

sztyn Ridge in the Pieniny Klippen Basin (PKB) during

the Jurassic, occurring in a 17 km long band in western

Slovakia, extending from the Dolná Súča Klippe on the SW,

through Krivoklát and Vršatec klippen till Mikušovce Klippe

on the NE (Fig. 1; Mišík 1979; Morycowa & Mišík 2005).

They form marked rocky cliffs and their thickness attains

several tens of meters. The carbonate sediment produced in

such environments during the Jurassic probably sourced deeper,

offshore and basinal environments of the Pieniny Klippen

Belt. However, the majority of the Jurassic sediments on

the Czorsztyn Ridge were deposited on pelagic, sediment-

starved seafloor at depths below the photic zone (Fig. 2), with

depositonal conditions that were similar to those occurring on

pelagic carbonate platforms during the Jurassic in the Eastern

Alps, Southern Alps, Apennines, and Sicily (Santantonio

1993; Cobianchi & Picotti 2001; Marino and Santantonio

2010). The biohermal limestones with corals define the so-called

Vršatec Limestone (Mišík 1979). Mišík (1979) and Morycowa

& Mišík (2005) suggested that the Vršatec Limestone belongs

to the Oxfordian stage on the basis of corals and bivalves.

Owing to this assignment of coral limestones to the Oxfordian,

Mišík (1979) distinguished two tectonic slices at the Vršatec

klippen. He suggested that these two slices significantly differ

in the development of Oxfordian deposits, the first one with

shallow water-coral biohermal limestones deposited in photic

environments, and the second one with strongly condensed,

red micritic nodular or non-nodular limestones deposited in

aphotic environments (Fig. 3). In contrast to Mišík (1979),

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Schlögl et al. (2006, 2009a, b) showed that the Vršatec Lime-

stone belongs to the Bajocian on the basis of the stratigraphic

superimposition criteria and ammonite occurrences. First,

the Middle Jurassic crinoidal limestones overlie coral lime-

stones at Vršatec-Javorníky Klippe, and coral limestones thus

cannot belong to the Oxfordian. Second, Upper Bajocian–

Lower Bathonian ammonite Nannolytoceras tripartitum occurs

in a dyke penetrating through the Vršatec Limestone (Schlögl

et al. 2006, 2009a). In addition, several other dykes with

Bathonian–Callovian ammonites occur in the uppermost part

of the Vršatec Limestone in the Vršatec-Castle Klippe (Schlögl

et al. 2009b). Crinoidal limestones in the Czorsztyn Unit in

the Polish and Slovakian parts of PKB (assigned to the Smole-

gowa and Krupianka formations) range from the upper Propin-

quans Zone of the Lower Bajocian to the lower Garantiana

Zone of the Upper Bajocian on the basis of ammonites (e.g.,

Krobicki & Wierzbowski 2004; Wierzbowski et al. 2004).

In the Czorsztyn Unit, these crinoidal limestones exhibit hia-

tuses both at their lower and upper boundaries.

Morycowa & Olszewska (2013) analyzed thin sections from

the biohermal and peribiohermal facies of the Vršatec Lime-

stone. They described benthic foraminifers of the genera

Ruma nolina, Paalzowella, Redmondoides, Troglotella, and,

Haghimashella and the microencruster Iberopora. They argued

that four foraminifer species and Iberopora appeared for

the first time in the Oxfordian, indicating that the Vršatec

Limestone is of Late Jurassic age, thus suggesting the initial

biostratigraphic inference of Mišík (1979) may be correct.

Here, using 120 new thin sections collected at five sites with

the Vršatec Limestone, we revisit stratigraphic and paleogeo-

graphic distribution of 14 species of benthic foraminifers from

the Vršatec Limestone, and assess overall genus-level compo-

sition of foraminiferal assemblages from several sites. We

show that the first appearances of species that were supposed

to be indicative of the Late Jurassic, as detected in other

Tethyan regions, actually extend to the Bajocian. We docu-

ment the presence of species that are alone or in combination

diagnostic of the Middle Jurassic, clarify stratigraphic and

geographic ranges of the identified species of benthic forami-

nifers, and discuss the stratigraphic position of the Vršatec

Limestone and its general biotic composition.

Geological and stratigraphic setting

The Pieniny Klippen Basin (Western Carpathians) was rep-

resented by a depositional belt of mixed, carbonate–siliciclas-

tic ramps with shallow-water and hemipelagic sedimentation.

Fig. 1. Geographic maps showing the location of the Pieniny Klippen Belt in the Western Carpathians and the locations of five sites of outcrops

for the Vršatec Limestone. A sixth site, the Krivoklat Gorge, was not sampled in this study and the occurrences near Mikušovce and Dolná Súča

(also not included) are out of the satellite map.

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This basin belonged to the sout hern

edge of the NW European platform

during the Early Jurassic and during

the Aalenian (Segit et al. 2015).

The PKB was located at 30–40 ºN

during the Middle Jurassic and at

20–30 ºN during the Late Jurassic

(Lewandowski et al. 2005). Although

some degree of syntectonic differen-

tiation into seamount-like elevations

and deeper basins took place already

during the Early Jurassic (Mišík et al.

1995), this belt became fully discon-

nected from the NW European plat-

form and differen tiated into shallower,

pelagic carbo nate platforms (Czor-

sztyn Ridge) and deeper basins

(Kysuca Basin) during the earliest

Bajocian (Birkenmajer 1977; Mišík

1979; Aubrecht 1997). The Bajocian–

Early Oxfordian sedimentation was

extremely sediment-starved and dis-

continuous on the Czorsztyn Ridge,

disconnected from river-born sediment

supply, and predominantly taking place

at aphotic depths with very limited in

situ carbonate production, with seve-

ral minor and major stratigraphic hia-

tuses (Fig. 2; Aubrecht & Szulc 2006;

Schlögl et al. 2009b).

The base of the Middle Jurassic is

characterized by the deposition of

hemipelagic dark marls and marl-

stones (Krempachy Marl, Skrzypne

Marl, and Harcygrund Marl forma-

tions). The termination of this marly

siliciclastic regime on the Czor sztyn

Ridge is marked by a hardground

with lag deposits on the base of cri-

noidal limestones of Bajocian age

(Smo le gowa and Krupianka for ma-

tions, Fig. 2). Schlögl et al. (2006)

sug gested that coral biohermal lime-

stones of the Vršatec Limestone deve-

loped on the shallowest portions of

the Czor sztyn Ridge prior to the depo-

sition of the crinoidal limestones or were time-equivalent with

crinoidal limestones of the Smolegowa and Krupianka forma-

tions. This view differs from initial assignment of the Vršatec

Limestone to the Oxfordian by Mišík (1979), indicating

that crinoidal limestones underlie coral limestones (Fig. 3).

The Vršatec Limestone is formed by coral biohermal framesto-

nes, bindstones, and rudstones. In addition to reef constructors,

benthic communities are dominated by bivalve assem blages.

Lime stones with coral reefs are horizontally and/or laterally

replaced by (i) breccias that accumulated at footwall margins

of faulted blocks (with clasts of biohermal limestones) and by

(ii) crinoidal limestones. After a hiatus (marked by hard-

ground), these biohermal limestones are almost always over-

lain by crinoidal– spiculitic limestones (Fig. 4).

Crinoidal limestones locally alternate or pass upwards into

stromatactis-rich limestones with frequent relicts of sponges

(Aubrecht et al. 2002, 2009). The spiculitic limestones were

deposited during the Bajocian in slope and basinal

environments of the Czorsztyn Ridge (Flaki and Podzamcze

formations, Birkenmajer 1977). The crinoidal limestones and

HIATUS

Czorsztyn Ridge

edge

platform

Aalenian

Bajocian

Bathonian

Callovian

Oxfordian

Kimmeridgian

Tithonian

Early

Late

Middle

Early

Middle

Fig. 2. The lithostratigraphic scheme of the Czorsztyn Ridge for the Middle and Upper Jurassic

along an edge-platform gradient, with the Vršatec Limestone deposited in the shallowest parts of

the Czorsztyn Ridge. The stage durations are proportional to durations in Ogg et al. (2016).

1 — Dark marly deposits, undivided Krempachy Marl Formation (lower part), and Skrzypny

Shale Formation (upper part). 2 — White to red crinoidal limestones, Smolegova and Krupianka

formations. 3 — Coral biohermal limestone and peri-biohermal limestone, Vršatec Limestone.

4 — Red non- nodular massive limestone, Bohunice Formation. 5 — Red nodular limestone,

Czosztyn Limestone Formation. 6 — Crinoidal–brachiopod limestone, Štepnice Limestone

Member. 7 — White-yellowish ammonite–brachiopod shell-beds, Kočkovce Limestone Member.

8 — White to red ammonite shell-beds, Rogoznik Coquina Member. 9 — Yellowish to reddish

massive bioclastic limestone, Korowa Limestone Member.

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their time-equivalents were deposited

during the Early Bajocian (late Pro-

pinquans and Humphriesianum zones)

and the early Late Bajocian (Niortense

Zone and the early part of the Garan-

tiana Zone). They are terminated by

another spatially extensive hardground,

with hiatus corresponding to the Late

Bajocian upper Garantiana and lower

Parkinsoni zones (Fig. 2; Wierzbowski

et al. 2004; Schlögl et al. 2005).

The hia tus between the cri noidal lime-

stones and the overlying formations is

present in all sections belonging to

the former Czorsztyn Ridge. The higher

parts of the Middle Jurassic suc-

cessions are represented by condensed

micritic to bioclas tic nodular Ammo-

nitico Rosso (Czor sztyn Formation)

or non-nodular mic ritic or bioclastic

limestones (Bohunice and Štepnica for-

mations) on the top of the Czor sztyn

Ridge and its slope (Mišík et al.

1994a; Aubrecht et al. 2009), and by

radiolarites and radio larian limestones

in the adjacent basins (Czajakowa

and Sokolica Radiolarite formations).

The Callo vian to Lower Oxfordian

depo

sits are rarely preser

ved on

the Czorsztyn Ridge and are mainly

represented by a major strati graphic

hiatus (Schlögl et al. 2009b).

Material and methods

The studied sites with biohermal–

peribiohermal limestones of the Vrša-

tec Limestone are situated in the wes tern

Slovakia, Middle Váh Valley, between

Vršatské Podhradie, Červený Kameň

and Krivo klát villages (Fig. 1). Coral-

domi nated bioherms are otherwise

absent in the rest of the Pieniny Klip pen Belt. They include:

(1) Vršatec Castle Klippe (VH, prima rily with peri-biohermal

facies with breccias, 49°03’55.57” N, 18°09’03.80” E),

(2) Vršatec-Javorníky Klippe (VJ, mainly with massive bio-

hermal facies, 49°04’10.24” N, 18°09’20.54” E); (3) Drieňová

Hora Klippe (DRIE, 49°02’26.44” N, 18°09’23.19” E), (4)

Malé Hradište Klippe (MH, 49°03’6.81” N, 18°11’31.24” E),

and (5) an unnamed klippe between Malé Hradište Hill and

Kalvária Hill (MH-K, 49°03’23.74” N, 18°11’39.03” E).

The Vršatec Limestone also occurs at four other sites (Kri vo-

klát Gorge, Mikušovce-quarry, Mikušovce-meadow, and

Mikušovce-Mn-mine) that were not investi gated in this study.

The Middle Jurassic-Lower Cretaceous limestone successions

at all localities belong to the Czorstyn Unit, and are capped by

the Upper Cretaceous marls.

Benthic foraminifers were studied in a total of 120 thin

sections prepared from samples collected from surface out-

crops (88 thin sections from two Vršatec Klippen, 17 thin sec-

tions from from Malé Hradište Klippe and MH-K Klippe, and

4 thin sections from Drieňová Hora Klippe). To document

differences between the lower and upper part of the formation,

multiple thin sections were collected at locality 22 (Mišík

1979, locality VJ 22 in this paper) and in a transect between

localities 24 and 40 (Mišík 1979, locality VJ 5 in this paper).

The microfossils in these samples were documented by

more than 3000 photographs. Zeiss microscope was used for

Fig. 3. The comparison of a lithostratigraphic succession at Vršatec–Javorníky interpreted in this

study (left column) and by Mišík (1979) (right column). Mišík (1979) invoked the presence of

two tectonic slices differing in the development of the Oxfordian deposits and in stratigraphical

polarity, which were separated by a thrust on the top of crinoidal limestones (dashed line).

However, in this study, we show that the klippe preserves a single Middle Jurassic–Lower

Cretaceous succession. Note: Lst. Fm. — Limestone Formation

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micropaleontological study of the thin sections and micropho-

tographs were taken with a Zeiss Axiocam 105 color digital

camera. Thin-sections are stored at the Department of Geology

and Paleontology, Faculty of Natural Sciences, Comenius

University in Bratislava (archived under the reference title of

this article). Some well-preserved specimens of Cornuspira,

Ophthalmidium, and Paalzowella were determined to species

level. All foraminifers that can be determined to genus level

were counted in thin sections. To summarize differences in

species abundances among the five sites and between the lower

and upper part of the formation, the counts were pooled to

site-level genus abundances and compared in barplots.

Results

Systematic paleontology

We use supraordinal classification of Foraminifera of

Pawlowski et al. (2013). The genus and species-level determi-

nation follows Loeblich & Tappan (1988, 1992), Clerc (2005),

and Rigaud et al. (2013, 2015a, b, 2018). The stratigraphic and

paleogeographic distribution of agglutinated benthic forami-

nifers of the order Lituolida is relatively well-documented

from the Jurassic deposits, especially on the basis of speci-

mens extracted from sieved unlithified samples (e.g., Tyszka

1994; Smoleń 2012). However, biostratigraphic and paleogeo-

graphic importance of small-sized species with calcareous

tests, belonging to the orders Miliolida, Spirillinida, and

Involutinida that frequently occur in lithified carbonate depo-

sits, remains poorly known. The representatives of the order

Miliolida possess an imperforate wall formed by high-Mg

calcite, with randomly oriented crystals refracting light in all

directions and resulting in a porcelaneous appearance of

the test. The representatives of the order Spirillinida generally

possessed low-Mg calcite, hyaline tests. The representatives

of order Involutinida have aragonitic tests.

Subphylum FORAMINIFERA d’Orbigny, 1826

Class TUBOTHALAMEA Pawlowski, Holzmann &

Tyszka, 2013

Order MILIOLIDA Delage & Hérouard, 1896

Suborder Miliolina Delage & Hérouard, 1896

Superfamily Cornuspiroidea Schultze, 1854

Family Cornuspiridae Schultze, 1854

Subfamily Cornuspirinae Schultze, 1854

Genus Cornuspira Schultze, 1854

Cornuspira infraoolithica Terquem, 1870

(Fig. 5.22–23)

1870 Cornuspira infraoolithica n.sp. – Terquem: p. 243, pl. XXV,

fig. 13.

2005 Cornuspira infraoolithica Terquem – Clerk: p. 36, pl. 3, fig.

1-4; pl. 14, fig. 6, 7.

Occurrences: Drieňová Hora Klippe (DRIE 01a), Malé

Hradište Klippe (MH 01/1).

Description: A small biconcave species with a test diameter

close to 250 μm, composed of a very small proloculus,

followed by 7 to 9 planispiral whorls.

Distribution: Cornuspira infraoolithica was described by

Terquem (1870) from the Upper Bajocian (Parkinsoni Zone)

of Moselle (France), by Terquem & Berthelin (1875) from

the Pliensbachian (Margaritatus Zone) of France, and by

Burbach (1886) from Pliensbachian of Gotha (Central

Germany). Clerc (2005) described this species from the Upper

Aalenian–Upper Bathonian deposits of the French Jura.

Stratigraphic range: Pliensbachian–Upper Bathonian.

Cornuspira orbicula (Terquem & Berthelin, 1875)

(Fig. 5.16–21)

1875 Cornuspira orbicula n. sp. – Terquem & Berthelin: p. 17, pl. I,

fig. 12a-c.

2005 Cornuspira orbicula (Terquem & Berthelin) – Clerc: p. 37, pl.

3, fig. 5-19; pl. 14, fig. 1-5; pl. 32, fig. 1, 2.

Occurrences: Vršatec Castle Klippe (VH 5B/2, VH 5B/2B),

Malé Hradište Klippe (MH 01/1, MH 01/3a, MH 01/6a,

MH 03/1a, MH 03/1c).

Description: A slightly biconcave medium-sized species

with a test diameter close to 380 μm, composed of a spherical

proloculus and a low tubular deuteroloculus followed by 5 to

6 planispiral whorls. This species is characterized by a high

morphological variation.

Discussion: Copestake & Johnson (2014) accepted that

Cornuspira liasina appears to be microspheric form of the spe-

cies while Terquem & Berthelin’s (1875) Spirillina orbiculare

seems to be its megalospheric equivalent.

Distribution: It is one of the most common representatives

of the genus Cornuspira in the Middle Jurassic. This species

was described as Spirillina orbicula by Terquem & Berthelin

(1875) from the Upper Pliensbachian (Margaritatus Zone) of

Essey-les-Nancy, France. In the first half of the 20

century,

it was reported from Lower and/or Middle Jurassic outcrops:

Lower Jurassic (Lias α, β, ζ) of the Swabian Alb (Issler 1908;

Franke 1936), Lower and Middle Jurassic of NW Germany

(Bartenstein & Brand 1937) and Callovian of NW Germany

(Lutze 1960). Antonova (1959) described this species from

the Lower Aalenian of the Laba River region (NW Caucasus).

Trifonova (1961) described this species from the Pliensbachian

and Toarcian of the villages of Sarantsi and Zimevitsa, Sofia

district (Bulgaria). C. orbicula is also known from the Lower

Toarcian of SW Germany (Riegraf 1985), and from the Aale-

nian–Lower Oxfordian (Wernli 1970) and Aalenian–Callovian

(Clerc 2005) of the French Jura.

Stratigraphic range: Lower Jurassic to Callovian, common

in the Upper Aaalenian, Upper Bajocian and Bathonian.

Cornuspira tubicomprimata Danitch, 1971

(Fig. 5.12–15)

1971 Cornuspira tubicomprimata sp. nov. – Danitch: p. 97, tabl. XVII,

fig. 2 а, b.

2005 Cornuspira tubicomprimata Danitch – Clerc: p. 40, pl. 3,

fig. 21-27; pl. 14, fig. 10, 11.

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Genus Meandrovoluta Fugagnoli & Rettori in Fugagnoli,

Giannetti & Rettori, 2003

Meandrovoluta asiagoensis Fugagnoli & Rettori, 2003

(Fig. 6.1–6)

1966 Glomospira sp. – Radoičić: pl. 92, fig. 2; pl. 111, fig. 2; pl.

124, fig. 1–2.

2003 Meandrovoluta asiagoensis gen. et sp. nov. – Fugagnoli &

Rettori: p. 45, pl. 1, figs. 1-12; pl. 2, figs. 1-5?, 6-16.

Occurrences: Vršatec Castle Klippe (VH 5B/1, VH 5B/2,

VH 5B/4, VH 5B/5a, VH 5B/5b, VH 5B/6, VH II/2, VH 6),

Drieňová Hora Klippe (DRIE 01, DRIE 02), Malé Hradište

Klippe (MH 01/6a, MH 03).

Description: This species shows a high morphological

varia tion. The coiling is zig-zag-shaped in the early stage and

slightly undulated (meander-like) in the later ontogenetic

stage. A globular proloculus is followed by an undivided

second chamber. The second chamber is mostly irregularly

coiled, with 3 to 5 coils. Most of the sections are identical to

the megalospheric forms of M. asiagoensis (e.g., figs. 1a, 2, 3,

5, 6–16 on pl. 2 in Fugagnoli et al. 2003) characterized by

the small size, by an increase of the zig-zag-shape part, and by

a reduction of the disc-like or fanlike second stage.

Distribution: A genus Meandrovoluta (type-species

Meandrovoluta asiagoensis) was originally described from

the Sinemurian?–Domerian interval of the Calcari Grigi

Formation of the Trento Platform (Southern Alps; Fugagnoli

et al. 2003). Abundant foraminifers similar to glomospirinids

occur in the Lower Jurassic limestone successions at many

locations of the Tethyan region. All these glomospirinid fora-

minifers occurring in the Lower Jurassic of the Tethyan region

were typically assigned to the genus Glomospira Rzehak,

1885 with a finely agglutinated wall (e.g., Radoičić 1966).

However, Meandrovoluta differs from Glomospira in having

a porcelaneous wall. Fugagnoli and Rettori (in Fugagnoli et

al. 2003) placed Meandrovoluta in the family Cornuspiridae

Schultze (order Miliolida) that is characterized by porcela-

neous tests. Cai et al. (2006) described three new species:

Glomo spira wölongensis, Glomospira tingriensis and Glomo

spirella minuscula from the Niehnieh Hsiungla Formation of

Tingri and Nyalam regions of southern Tibet, China. We

assume that the species Glomospira wölongensis (pl. I, figs.

1–10 in Cai et al. 2006) and Glomospirella minuscula (pl. I,

figs. 11–18 in Cai et al. 2006) represent junior syno nyms of

Meandrovoluta asiagoensis, because of the small size, very

thin wall, the mode of coiling of the second chamber, and

the poorly developed second disc-like stage. Based on ammo-

nites, the age of the formation is Bajocian to Callovian (Dhital

2015), but the ammonites come from other sections, not from

those studied by Cai et al. (2006). Therefore, the last occur-

rence date of this species on the basis of its distribution in

the Niehnieh Hsiungla Formation is poorly constrained.

M. asiagoensis was reported from the Sinemurian–Toarcian of

the Karst Dinarides, Croatia (Velić 2007), the Upper Sine-

murian–Upper Toarcian of Latium–Abruzzi carbonate plat-

form, Central Italy (Chiocchini et al. 2008), and the Upper

Sinemurian–Lower Pliensbachain of the Podpec Limestone of

External Dinarides, Slovenia (Gale 2014).

Stratigraphic range: Sinemurian–Toarcian, Middle Jurassic.

Superfamily Milioloidea Ehrenberg, 1839

Family Spiroloculinidae Wiesner, 1920

Genus Labalina Azbel, 1988

Labalina occulta (Antonova, 1958)

(Fig. 7.1–4)

1958 Spirophthalmidium occultum sp. n. – Antonova: p. 52, tabl. II,

fig. 5а, b, 6.

2005 Labalina occulta (Antonova) – Clerc: p. 78, pl. 11, fig. 10-22;

pl. 26, fig. 4-15.

Occurrences: Vršatec Castle Klippe (VH 5B/3, VH 5B/4),

Malé Hradište Klippe (MH 01/1, MH 03).

Description: The longitudinal sections have an oval contour.

The length of the tests is around 200 μm. The tests are charac-

terized by well-rounded evolute arrangement of the chambers

and by a thin wall. In transverse sections, the specimens are

mainly oval and stretched, with well-visible coiling in a quin-

queloculinid arrangement, followed by a sigmoidal arrange-

ment and a lobed contour. The number of chambers varies

between 9 and 12.

Distribution: Antonova (1958) described this species as

Paleomiliolina occulta from the Bajocian of the Psebai district,

NW Caucasus, and from the Bajocian of the Laba River area,

NW Caucasus (Antonova 1959). Danitch (1971) reported this

species from the Upper Bajocian–Middle Bathonian of

the region between Dniester and Prout rivers, Moldavia. Clerc

(2005) found it in the Middle Aalenian to Upper Bathonian of

the French Jura.

Stratigraphic range: Middle Aalenian–Upper Bathonian.

Superfamily Nubecularioidea Jones, 1875

Family Nubeculariidae Jones, 1875

Subfamily Nubeculariinae Jones, 1875

Genus Nubecularia Defrance, 1825

Nubecularia reicheli Rat, 1966

(Fig. 5.7–11)

1966 Nubecularia reicheli n. sp. – Rat: p. 80, fig. 2; pl. 1, fig. 1-9.

2005 Nubecularia reicheli Rat – Clerc: p. 52, pl. 17, fig. 7-9; pl. 18,

fig. 1-6.

Occurrences: Vršatec Castle Klippe (VH 5B/1, VH 5B/2,

VH 5B/3, VH 5B/4, VH 5B/5a, VH 5B/5b, VH 5B/6, VH 5B/7,

VH 5B/8, VH II/1, VH II/2a, VH II/2b, VH II/4, VH Bos

Lms/B, VJ 22, VH 100m, and samples from “Bositra dyke”

(VH-Bositra B). Vršatec-Javorníky Klippe (VJ 5/2c, VJ 5/top),

Drieňová Hora Klippe (DRIE 01), Malé Hradište Klippe

(MH 01/1, MH 01/6a, MH 01/7, MH 02/1, MH 02/2b,

MH 02/4a, MH 02/5, MH 03, MH 03/1a, MH 03/1c,

MH 01/new, MH 02/new, MH 03/new, MH GPS).

Description: Encrusting porcelaneous species, characte-

rized by thick crusts consisting of several layers and forming

millimetric platoons.

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Distribution: This species was described by Rat (1966)

from the Bajocian of Burgundy (NE France), and by Wernli

(1970) and Clerc (2005) from the Lower Bajocian to

the Middle Callovian and the Upper Bajocian to the Lower

Callovian of the French Jura, respectively.

Stratigraphic range: Lower Bajocian–Middle Callovian.

Subfamily Nubeculinellinae Avnimelech & Reiss, 1954

Genus Vinelloidea Canu, 1913

Vinelloidea bigoti (Cushman, 1930)

(Fig. 5.1–6)

1930 Nubeculinella bigoti n. sp. – Cushman: p. 134, pl. IV, fig. 2, 3.

2005 Nubeculinella aff. bigoti Cushman – Clerc: p. 47, pl. 15, fig. 1,

2, 5-8.

Occurrences: Vršatec Castle Klippe (VH 5B/1, VH 5B/2,

VH 5B/3, VH 5B/4, VH 5B/5a, VH 5B/7, VH II/1, VH II/2b,

VH 100m, Vršatec-Javorníky Klippe (VJ 5/2c), Malé Hradište

Klippe (MH 01/1, MH 02/4a, MH 02/5, MH 03, MH 03/1a).

Description: The subsphaerical proloculus is followed by

a narrow, tube-like chamber, up to five chambers in the initial

whorl are visible and up to four chambers in the uncoiled,

linear portion of the test. The wall is calcareous, built by

high-Mg calcite.

Remarks: Canu (1913) assigned the genus Vinelloidea

(type species V. crussolensis) to bryozoans. Cushman (1930)

introduced the genus Nubeculinella from the Jurassic of

Auberville (Calvados, France), with type species Nubeculinella

bigoti. Voigt (1973) showed that the genus Vinelloidea Canu

represents an adherent foraminifer, and is a senior synonym of

the genus Nubeculinella Cushman. Loeblich & Tappan (1988)

retained the genus name Vinelloidea Canu and refigured

the type material of V. crussolensis Canu (Voigt 1973), desig-

nating it as a lectotype. Copestake & Johnson (2014) sugges-

ted that illustrations of Vinelloidea crussolensis in Loeblich &

Tappan (1988) and Nubeculinella bigoti in Loeblich & Tappan

(1988) do not warrant species discrimination and thus syno-

nymized the two species. We follow this suggestion and use

the genus name Vinelloidea on the basis of the priority rule.

Distribution: The species was first described by Cushman

(1930) as Nubeculinella bigoti from the Oxfordian (Cardio

ceras cordatum Zone) of Auberville (Calvados, France).

Wide spread in the Tethyan region, it was found by Paalzow

(1932) in the Middle Oxfordian (Trans versarium Zone) of

NE Swabian Alb, S Germany, by Gordon (1961, 1965) in

the Oxfordian–Kimmeridgian of Dorset (S. England), by

Antonova (1959) in the Aalenian–Lower Bajocian of the Laba

River region, Russia, by Adams (1962) in the Lower Jurassic

to Kimmeridgian of England, by Danitch (1971) in the Upper

Oxfordian–Lower Kimmeridgian of the region between

Dniester and Prut rivers, Moldavia. The species was described

from the Toarcian–Oxfordian of the French Jura, France by

Clerc (2005) and from the Lower Jurassic (Obtusum to

Jamesoni zones) of Llanbedr (Mochras Farm) Borehole, North

Wales, UK by Copestake & Johnson (2014).

Stratigraphic range: Sinemurian to Kimmeridgian.

Family Ophthalmidiidae Wiesner, 1920

Genus Ophthalmidium Kubler & Zwingli, 1870

Ophthalmidium caucasicum (Antonova, 1958)

(Fig. 7.15–19)

1958 Spirophthalmidium caucasicum sp. n. – Antonova: p. 51, tabl.

II, fig. 1-4.

2005 Ophthalmidium caucasicum (Antonova) – Clerc: p. 55, pl. 4,

fig. 3-8; pl. 20, fig. 1-10; pl. 32, fig. 3, 4.

Occurrences: Malé Hradište Klippe (MH 01/new, MH 03/

new).

Description: A rather large species (up to 440 μm) charac-

terized by a fully evolute test with a rounded peripheral mar-

gin, the wall is fine, markedly thin (not exceeding 10 μm).

Distribution: The species was first described by Antonova

(1958) as Spirophthalmidium caucasicum from the Bajocian

of the Psebai district, NW Caucasus. Antonova (1959)

documented this species also from the Bajocian of the Laba

River area (NW Caucasus) and Danitch (1971) from the Upper

Bajocian–Lower Bathonian of the region between Dniester

and Prut rivers, Moldavia. It was described by Clerc (2005)

from the Bajocian–Lower Bathonian of the French Jura.

Stratigraphic range: Bajocian–Lower Bathonian.

Ophthalmidium obscurum (Ivanova & Danitch, 1971)

(Fig. 7.20–29)

1971 Spirophthalmidium obscurum sp. n. – Ivanova & Danitch in

Danitch: p. 126, tabl. XXVIII, fig. 1а, b, 2а, b, 3, 4.

2005 Ophthalmidium obscurum (Ivanova & Danitch) – Clerc: p. 69,

p. 7, fig. 8-14; pl. 22, fig. 6-10.

Occurrences: Vršatec Castle Klippe (VH 5B/1, VH 5B/2,

VH 5B/3, VH 5B/4, VH 5B/6, VH 5B/7, VH 5B/8, VH II 2b,

VH II 4, VH 100m, and samples from „Bositra dyke“

(VH-Bositra A). Vršatec-Javorníky Klippe (VJ 5t, VJ22), Malé

Hradište Klippe (MH 01/1, MH 02/2b, MH 03/1a).

Description: A medium-sized species (up to 340 μm)

characterized by an involute coiled test, oval shape, and a wall

thick up to 18 μm.

Distribution: The species was first described by Danitch

(1971) from the Upper Bajocian–Lower Bathonian of

the region between Dniester and Prout rivers (Moldavia).

Wernli (1970) described the species from the Upper Bajocian

of the French Jura and Clerc (2005) from the Upper Bajocian–

Lower Bathonian of the French Jura.

Stratigraphic range: Upper Bajocian to Lower Bathonian.

Ophthalmidium terquemi Pazdrowa, 1958

(Fig. 7.5–14)

1958 Ophthalmidium carinatum n. subsp. terquemi – Pazdrowa: p.

152, tabl. I, fig. 1-9; tabl. II, fig. 11; tabl. III, fig. 1-8; tabl. V,

fig. 7; tabl. VI, fig. 1-3; tabl. VII, fig. 5-7.

2005 Ophthalmidium terquemi Pazdrowa – Clerc: p. 71, pl. 7, fig.

17-21; pl. 8, fig. 1-17; pl. 9, fig. 1-3; pl. 23, fig. 2-9; pl. 33,

fig. 1, 2.

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Occurrences: Vršatec-Javorníky Klippe (VJ 10base), Drie-

ňová Hora Klippe (DRIE 01), Malé Hradište Klippe (MH 01/6b,

MH 02/2b, MH 03, MH 03/1a).

Description: Relatively large species (up to 500 μm)

characterized by an involute to semi-involute coiled test and

with a relatively thick wall (12 and 30 μm) for the last cham-

ber and the inner wall of the chambers is thicker at the base

and at the collar level.

Distribution: The species Ophthalmidium terquemi

Pazdrowa was described by Pazdrowa (1958) as a new sub-

species of Ophthalmidium carinatum Kubler & Zwingli from

the Bajocian of Czestochowa (Poland). This taxon was distin-

guished as Ophthalmidium carinatum terquemi Pazdrowa

from the Bajocian–Middle Bathonian of Czestochowa, Poland

(Pazdrowa 1959; Pazdro 1972), and as Ophthalmidium terquemi

from the Upper Bajocian–Lower Callovian of the Swabian

Alb (Blank 1990). It was found in the Bajocian–Lower

Bathonian of the French Jura and Burgundy by Piuz (2004)

and in the Bajocian–Lower Callovian of the French Jura by

Clerc (2005).

Stratigraphic range: Bajocian–Lower Callovian.

Order SPIRILLINIDA Gorbatchik & Mantsurova, 1980

Suborder Spirillinina Hohenegger & Piller, 1975

Family Spirillinidae Reuss & Fritsch, 1861

Subfamily Neotrocholininae Rigaud, Schlagintweit & Bucur,

2018

Genus Hungarillina Blau & Wernli, 1999

Rigaud et al. (2018) comprehensively revised this genus,

with synonymy lists and new taxonomic, phylogenetic, and

stratigraphic schemes, and assigned it to the subfamily

Neotrocholininae.

Hungarillina lokutiense Blau & Wernli, 1999

(Fig. 6.7–11)

1999 Hungarillina lokutiense n. gen., n. sp. – Blau & Wernli, p. 539,

pl. I, figs. 1-17.

2018 Hungarillina lokutiense Blau & Wernli – Rigaud,

Schlagintweit & Bucur, figs. 2F-G.

Occurrences: Vršatec Castle Klippe (VH 5B/2, VH 5B/5b),

Malé Hradište Klippe (MH 03).

Description: Relatively high trochospirally coiled test with

completely filled umbilical cavity displays the characteristic

“bell shape” of the type-species.

Distribution: Blau & Wernli (1999) described their new

genus Hungarillina with three new species: H. lokutiense,

H. media and H. pedunculata from the Middle Jurassic pebbles

in the Upper Bajocian megabreccia near Lokut (Transdanubian

Central Range, Hungary). Velledits & Blau (2003) described

this species from protoglobigerinid wackestones and pack-

stones from neptunian dykes in the Büdöskút Limestone,

Bükk Mountains (NE Hungary). Therefore, the stratigraphic

range of this species can extend beyond the Bajocian stage.

This species was also found by Piuz (2004) in the Lower

Bajocian of the French Jura and in Burgundy (SE France).

Schlagintweit & Moshammer (2015) described this species

from a fissure (filled by peloidal packstones and grainstones)

in the Vils Limestone (Außerfern in Tyrol, Austria). Its Bajo-

cian age was inferred on the basis of H. lokutiense.

Discussion: The microfacies with protoglobigerinids and

spirillinids, containing Hungarillina lokutiense and Radio

spirillina umbonata, was assigned by Velledits & Blau (2003)

to the Bathonian?–Callovian?.

Stratigraphic range: Bajocian (this paper), Bathonian?–

Callovian? (Velledits & Blau 2003).

Genus Radiospirillina Blau & Wernli, 1999

Rigaud et al. (2018) assigned this genus to subfamily

Neotrocholininae.

Radiospirillina umbonata Blau & Wernli, 1999

(Fig. 6.27–30)

1999 Radiospirillina umbonata n. gen., n. sp. – Blau & Wernli: p.

541, pl. II, figs. 3, 5, 7-8.

2018 Radiospirillina umbonata Blau & Wernli – Rigaud,

Schlagintweit & Bucur: fig. 4A.

Occurrences: Vršatec Castle Klippe (VH 5B/1, VH 5B/2,

VH 5B/5a), Malé Hradište Klippe (MH 03/1a, MH 03).

Description: Sublenticular test, formed by a globular pro-

loculus followed by subplanispirally to low trochospirally

enrolled, undivided second tubular chamber.

Distribution: The genus Radiospirillina, with the type spe-

cies Radiospirillina umbonata, was described as a new genus

and a new species by Blau & Wernli (1999) from the Middle

Jurassic pebbles in the Upper Bajocian megabreccia near

Lokut (Transdanubian Central Range, Hungary). Velledits &

Blau (2003) described this species from neptunian dykes in

the Büdöskút limestone, Bükk Mountains (NE Hungary). Piuz

(2004) reported the same species from the Bajocian echino-

dermic shelf in the French Jura and in Burgundy (SE France).

A specimen illustrated by Mišík et al. (1994b, pl. 2, fig. 6) and

determined as Trocholina sp., found in a dyke penetrating

through Bajocian pink crinoidal limestones (formerly sup-

posed to be of Bathonian-Callovian age) from the Krasín

Klippe near Dolná Suča also belongs to Radiospirillina

umbonata.

Stratigraphic range: Bajocian, Bathonian?– Callovian?

(Velle dits & Blau 2003).

Genus Tethysiella Blau, 1987a

Tethysiella pilleri (Blau, 1987a)

(Fig. 6.12–21)

1987a Praepatellina pilleri n. gen. n. sp. – Blau: p. 508, pl. 3, figs.

9-11, 13-15.

1991 Tethysiella pilleri (Blau, 1987a) – Blau & Haas: p. 21, figs. 7

S-T.

Occurrences: Vršatec Castle Klippe (VH 5B/2, VH 5B/5b),

Vršatec-Javorníky Klippe (VJ 5/2c), Malé Hradište Klippe

(MH 01/3a, MH 03, MH 03/1a, MH 03/1c).

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Description: Trochospirally coiled test consists of two

chambers, proloculus and a very high trochospiral deutero-

loculus tube, and possessing an empty umbilicus.

Distribution: Tethysiella pilleri was described as

Praepatellina pilleri by Blau (1987a) from the red fissure

fillings in the Oberrhät Limestone (Lavanter Breccie, Lienzer

Dolomiten, Austria, East-Tyrol). The fissure fillings are of

the Early Jurassic age. Blau & Haas (1991) found this species

in red fissure fillings in the Lower Jurassic limestones from

Transdanubian Central Range (Hungary). They changed

the genus name of this species, characterized by a very high

trochospiral deuteroloculus tube and an empty umbilicus, to

Tethysiella Blau, 1987a (Praepatellina Blau, 1987a is a junior

homonym of Tethysiella). The species was reported also from

the Bajocian of the French Jura and Burgundy, SE France

(Piuz 2004, 2008).

A specimen illustrated by Mišík et al. (1994b, pl. 2, fig. 5)

and determined as Trocholina sp., found in a dyke penetrating

through Bajocian pink crinoidal limestones (formerly assigned

to Bathonian–Callovian) from the Krasín Klippe near Dolná

Súča also belongs to Tethysiella pilleri.

Stratigraphic range: Lower Jurassic–Bajocian.

Family Placentulinidae Kasimova, Poroshina &

Geodakchan, 1980

Subfamily Ashbrookiinae Loeblich & Tappan, 1984

Genus Paalzowella Cushman, 1933

Paalzowella turbinella (Gümbel, 1862)

(Fig. 8.1, 2)

1862 Rotalina turbinella n. sp. – Gümbel: p. 230, taf. IV.

fig. 10a-b.

2015 Paalzowella? sp. aff. turbinella (Gümbel) – Schlagintweit &

Moshammer: p. 211, text-figs. 3 pars, 4a-f.

Occurrences: Vršatec-Javorníky Klippe (VJ 5/2a, V11ts),

Malé Hradište Klippe (MH 01/1, MH 01/3a)

Description: We assign the trochospirally coiled specimens

of the genus Paalzowella whith convex umbilical side dis-

playing a central flattened part and giving rise to a boat-like

outline of the test in axial sections to P. turbinella (in accor-

dance with the diagnostic trait of the species).

Distribution: Gümbel (1862) introduced the new species

Rotalina turbinella from the Middle Oxfordian at Streitberg

(Franconian Alb). A specimen illustrated by Mišík et al.

(1994b, pl. 2, fig. 7) and determined as Schakoinella cf.

spinata Blau, found in a dyke penetrating through Bajocian

pink crinoidal limestones (formerly supposed to be of

Bathonian–Callovian age) from the Krasín Klippe near Dolná

Súča also belongs to Paalzowella turbinella. The species was

described from the Bathonian of the Mecsek Mountains (South

Hungary) by Görög (1995). Schlagintweit & Moshammer

(2015) found the species in a fissure filling in the Vils

Limestone (Eastern Alps) of Bajocian age (age determination

on the basis of presence of H. lokutiense). Morycowa &

Olszewska (2013) described this species from the Vršatec

Limestone.

Stratigraphic range: Bajocian–Middle Oxfordian.

Paalzowella feifeli (Paalzow, 1932)

(Fig. 8.3–23)

1932 Trocholina feifeli nov. spec. – Paalzow: p. 140, taf. XI, fig. 6, 7.

1932 Trocholina elevata nov. spec. – Paalzow: p. 140, taf. XI, fig. 4.

1932 Trocholina transversarii nov. spec. – Paalzow: p. 141, taf. XI,

fig. 8, 9, 10.

2004 Paalzowella feifeli (Paalzow) – Piuz: PI. 4, figs 1-12, fig. 13-18,

P. feifeli ? formes carénées, fig. 15, 17, 18, P. feifeli aff. elevata

(Paalzow)

Occurrences: Vršatec Castle Klippe (VH 5B/5b), Vršatec-

Javorníky Klippe (VJ 5/1, VJ 5/A), Malé Hradište Klippe

(MH 01/1, MH 01/3a, MH 01/6a, MH 01/7, MH 02/1,

MH 02/2b, MH 03, MH 03/1a, MH 03/1c, MH 01/new,

MH 03/new, MH GPS).

Description: Trochospirally coiled in a low cone test

(Fig. 8.3–6 — Paalzowella feifeli feifeli) or in a high cone test

(Fig. 8.7–15 — Paalzowella feifeli elevata), numerous cham-

bers arranged in 5 to 12 whorls, periphery of the chambers

ornamented by carinae with elevated flangelike keel. In

the studied material we also found sections with a strongly

curved central part on the convex umbilical side and with very

long and strongly flangelike keels (Fig. 8.16–23, determined

here as Paalzowella sp.).

Distribution: Paalzow (1932) described three new species

of the genus Trocholina Paalzow, 1922: Trocholina feifeli,

T. elevata and T. transversarii from the Middle Oxfordian of

the Franconian Alb (Lower Schwammergel, Streitberg). As

noted by Schlagintweit & Moshammer (2015), most species

and subspecies of the genus Paalzowella (type species

Paalzowella turbinella (Gümbel, 1862)) were described from

the Middle to Upper Jurassic on the basis of isolated speci-

mens: by Seibold & Seibold (1960) (Oxfordian–Lower

Kimmeridgian of South Germany); Lutze (1960) (Lower

Oxfordian of Northwestern Germany); Bielecka (1960)

(Oxfordian of Chrzanow, Southern Poland); Bastien & Sigal

(1962) (Upper Oxfordian of Trept, Isere); Oesterle (1968)

(Oxfordian of the Swiss Jura Mountain); Winter (1970)

(Lower Kimmeridgian of Franke, Germany); Stam (1986)

(Lower Callovian–Upper Oxfordian of Portugal), Schmalzriedt

(1991) (Oxfordian–Lower Kimmeridgian of Swabian Alb,

SW Germany), Canales et al. (1993) (Upper Aalenian–

Lower Bajocian of the Southwest sector of the Basque-

Cantabrian basin, Spain), Görög (1995) (Bathonian of

the Mecsek Mountains, South Hungary), Görög et al. (2012)

(Callovian of the Villány Mountains, southern Hungary), and

others. Piuz (2004) documented this species in thin sections

from the Bajocian of the French Jura and Burgundy (SE

France). Although Morycowa & Olszewska (2013) argued

that the presence of Paalzowella turbinella and Rumanolina

feifeli (here assigned to the genus Paalzowella) supported

the Late Jurassic age of the Vršatec Limestone, these species

clearly originated earlier and are not diagnostic of the Late

Jurassic.

Stratigraphic range: Upper Aalenian–Lower Kim meridgian.

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Class FORAMINIFERA INCERTAE SEDIS

Order INVOLUTINIDA Hohenegger & Piller, 1977

Family Trocholinidae Kristan-Tollmann, 1963,

emend. Rigaud et al., 2013

Subfamily Lamelliconinae Zaninetti et al., 1987,

emend. Rigaud et al., 2013

Genus Kristantollmanna Rigaud, Blau, Martini & Rettori, 2013

Kristantollmanna cf. altissima (Pirini, 1966)

(Fig. 6.22, 23)

1966 Turrispirillina altissima n. sp. – Pirini: p. 95, taf. 3, fig. 1-3, ?4-5.

1987a Turrispirillina (?) altissima Pirini – Blau: S. 505, taf. 4,

fig. 10-13.

Occurrences: Drieňová Hora Klippe (DRIE 01, DRIE 06),

Malé Hradište Klippe (MH 03).

Description: High conical test consisting of globular pro-

loculus followed by a trochospirally enrolled, undivided tubu-

lar chamber, characterised by reduced lamellae on both sides

of the test.

Distribution: The species was described by Pirini (1966) as

Turrispirillina altissima from the Lower Jurassic limestones

of Montemerano-Grosseto (Central Italy). Blau (1987a) des-

cribed the species from the red fissure fillings in the Oberrhät

Limestone (Lavanter Breccie, Lienzer Dolomiten, Austria).

The fissure fillings were deposited during the Early Jurassic.

Stratigraphic range: Lower Jurassic, Bajocian (this paper).

Genus Trocholina Paalzow, 1922

Trocholina turris Frentzen, 1941

(Fig. 6.24, 25)

1941 Trocholina turris n. sp. – Frentzen: p. 306, taf. 1, fig. 13 a-c.

1999 Trocholina turris Frentzen – Böhm: p. 181, pl. 18, figs. 1-3.

Occurrences: Malé Hradište Klippe (MH 01/3, MH 01/7,

MH 03/1c).

Description: Conical test characterised by a high trocho-

spiral coiling of the deuteroloculus, the number of whorls is

7 to 10.

Distribution: The species was described by Frentzen (1941)

from the Lower Jurassic of SW Germany. It is known mainly

from the European epicontinental Lower Jurassic but also

from the Upper Triassic. Kristan-Tollmann (1990) described

this species from the Rhaetian of Central Papua New Guinea

and Senowbari-Daryan et al. (2010) from the Rhaetian of

the Nayband Formation (Central Iran). Blau (1987b) reported

T. turris from red fissure fillings in the Oberrhät Limestone

(Lavanter Breccie, Lienzer Dolomiten, Austria). Blau & Haas

(1991) described this species from red fissure infillings (Lower

Jurassic from Transdanubian Central Range, Hungary). Böhm

et al. (1999) depicted this species from the Hettangian–

Sinemurian of Adnet (Salzburg, Austria). Velledits & Blau

(2003) reported T. turris from crinoidal wackestones–pack-

stones in neptunian dykes in the Büdöskút Olistolith, Bükk

Mountains (NE Hungary). According to Velledits & Blau

(2003), the age of these crinoidal wackestones–packstones can

range from Hettangian to Sinemurian. However, their record

of the species is based on a single specimen only.

Stratigraphic range: Norian?, Rhaetian–Lower Jurassic,

Bajocian (this paper).

Discussion

Macrobenthic assemblages

Coral assemblages of the Vršatec Limestone were described

by Morycowa & Mišík (2005). We re-assessed the composi-

tion of the coral assemblages on the basis of new and exten-

sive sampling (Schlögl et al. 2014). The most abundant genera

are represented by Isastrea, Periseris, Thecosmilia,

Cladophyllia, Dendraraea, and Thamnasteria. Such coral

assemblage is typical of the Lower Bajocian reefs of France,

Luxembourg and Switzerland (Lathuilière 2000a, b). Five of

these genera are also common in the Oxfordian, especially at

higher-latitude reefs, but they are represented by morpholo-

gically similar but different species in the Bajocian and in

the Oxfordian. The genus Periseris does not occur in

the Upper Jurassic. Morycowa & Mišík (2005) described

the genus Atelophyllia on the basis of two fragments. We con-

firm the identification of this genus, which was known

from the Lower Bajocian of France only. The findings of

Dendraraea dendroidea are also consistent with the Bajocian

age (Lathuilière & Gill 1998). The Bajocian age also explains

the absence of some coral taxa that are generally very abun-

dant in Oxfordian reefs. Coral reefs were widespread on

the epicontinental shelves on the northern margin of the Tethys

Ocean during the Oxfordian (Insalaco et al. 1997; Leinfelder

et al. 2002; Martin-Garin et al. 2012). They formed also exten-

sive deposits on shallow platform margins in the Tethys

Ocean, today exposed in the Southern Alps, Slovenia, Croatia,

Albania, and Montenegro (Turnšek et al. 1981; Bosellini et al.

1981; Winterer & Bosellini 1981; Sartorio 1989). However,

Oxfordian deposits in the Penninic Ocean mark the maximum

relative sea level rise, recorded by maximum condensation on

shallow elevations and by maximum extent of deposition of

radiolarites in troughs. Oxfordian coral reefs were not recorded

from pelagic platforms in the Penninic Ocean. In this light,

their presence in the Oxfordian and their absence on pelagic

carbonate platforms during time intervals with shallower con-

ditions during the Middle and latest Jurassic was enigmatic.

Our study thus explains this paradox by showing that coral

reefs were formed in the Pieniny Klippen Belt during

the Bajocian and not during the Oxfordian.

Bivalves are mostly represented by internal molds; recrys-

tallized shells with preserved external surface are rare. They

are frequent in coral framestones, floatstones and rudstones

at Vršatec-Javorníky (locality 22 in Mišík 1979), including

Chlamys (Chlamys) textoria, Camptonectes (Camptonectes) sp.,

Spondylopecten (Spondylopecten) cardinatus, Plagiostoma

premutabilis, Pseudolimea cf. duplicata, „Placunopsis“ sp.,

Liostrea sp., Actinostreon gregareum, and Pinna sp.

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Spondylopecten (Spondylopecten) cardinatus is typical of coral-

reef habitats. Corals themselves are occasionally bioeroded by

bivalves. The ichnofossil Gastrochaenolites found in corals

most probably refers to the bivalve taxon Lithophaga.

Kochanová (1979) described 15 bivalve taxa from the Vršatec

Limestone. She distinguished several species of the genus

Chlamys, but they all belong to Chlamys (C.) textoria that is

characterized by high intraspecific morphologic variation

(Johnson 1984). With the exception of Chlamys (C.) cf. sub

textoria (Kochanová 1979), individuals of the genus Chlamys

found in the Vršatec Limestone belong to coarsely-ribbed and

intermediate morphotypes of Chlamys (C.) textoria of Johnson

(1984). Such morphotypes inhabited coral and sponge reefs in

other geographic regions (Johnson 1984). Mass occurrences

of large-sized valves of Oxytoma (Oxytoma) inaequivalvis

directly on corals or between the coral colonies were docu-

mented at Drieňová Hora Klippe. This bivalve association is

highly similar to that described in the Bajocian of the southern

French Jura (Lathuilière 1982).

Coral framestones and floatstones contain brachiopod

assemblages primarily formed by small-sized rhynchonellids

(Siblík 1979). They resemble Parvirhynchia-dominated

assemblages that are typical of Bajocian coral reefs (Almeras

& Lathuilière 1984). Red crinoidal limestones of the Krupianka

Formation, immediately overlying coral bioherms at Vršatec-

Javorníky Klippe, about 1 m above the top of the Vršatec

Limestone, contain brachiopod assemblages with

Capillirhynchia jaccardi, Morrisithyris phillipsiana,

Acanthothiris spinosa, Antiptychina haasi, Monsardithyris

ventricosa, and Striirhynchia subechinata. Peribiohermal

limestones, formed by breccias with clasts of crinoidal and

biohermal limestones and radiaxial cements, probably formed

on the margin of bioherms (after the shutdown of coral pro-

duction), contain fissures filled with shell concentrations with

Bositra buchi and with abundant brachiopods at Vršatec

-Castle. They also contain ammonite Nannolytoceras tripar

titum, the stratigraphic range of the species is from the Upper

Bajocian Parkinsoni Zone to Lower Bathonian Progracilis

Zone. Brachiopods are represented by Ferrithyris antiplecta,

Antiptychina bivallata, Monsardithyris uniplicata, Caucasella

rectecostata, and Parvirhynchia mutans. A similar assemblage

occurs in the uppermost parts of crinoidal limestones at

Slavnické Podhorie (Pevný 1969; Aubrecht et al. 2002) and at

Babiná (Mišík et al. 1994a), indicating that the fissure was

filled not later than during the Late Bajocian.

In addition to abundant crinoids, biohermal and especially peri-

biohermal limestones contain at some levels also gastropods,

decapods, echinoids, and holothurians. Cidaroid spines can be

also locally abundant, forming peculiar crinoidal–cidaroid-rich

limestones at Mikušovce. Ammonites and nautiloids are extre-

mely rare, always fragmented and very poorly preserved.

Stratigraphic distribution of foraminiferal assemblages

The micropaleontological analysis of thin sections of bio-

hermal and peribiohermal Vršatec Limestone show that

assemblages of benthic foraminifers in the Vršatec Limestone

consist of 32 genera (Table 1), with three species of Cornu

spira, three species of Ophthalmidium and two species of

Paalzowella. They contain taxa of the class Tubothalamea

(orders Miliolida and Spirillinida), Globothalamea (orders

Robertinida, Rotaliida and Textulariida), as well as Incertae

sedis orders Lagenida and Involutinida. In accordance with

Morycowa & Olszewska (2013), we document the presence of

Paalzowella feifeli and Paalzowella turbinella. Morycowa &

Olszewska (2013) also described Rumanolina seiboldi, Ruma

nolina elevata, Troglotella incrustans, and Haghimashella cf.

arcuata. However, the specimens documented on their figs.

4.4 and 4.5 do not belong to the genus Rumanolina introduced

by Neagu & Cirnaru (2001) because they do not show a diag-

nostic trait of the genus, i.e., acute to flap-like keels that

should be developed along the arched sutures. In addition,

Table 1: The list of genera of the Vršatec Limestone determined at

four sites, with total number of specimens and total number of thin

sections with at least one specimen.

ršatec-Castle

ršatec-Javorníky

Malé Hradište

Drieňová Hora

ršatec-Castle

ršatec-Javorníky

Malé Hradište

Drieňová Hora

Number of specimens

Number of thin sections

Cornuspira

Meandrovoluta

Labalina

Nubecularia

Vinelloidea

Ophthalmidium

Hungarillina

Radiospirillina

Tethysiella

Paalzowella

Kristantollmanna

Trocholina

Frondicularia

Lingulina

Nodosaria

Lenticulina

Astacolis

Marginulina

Reinholdella

Epistomina

Ammobaculites

Callorbis

Trochammina

Duotaxis

Verneulinoides

Reophax

Mesoendothyra

Redmondoides

Bigenerina

Textularia

Troglotella

Earlandia

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the description of the genus Rumanolina is based on few iso-

lated specimens and both R. seiboldi and R. elevata are proba-

bly restricted to the Valanginian. The specimens determined as

Spirillina sp. (figs. 4.4 and 4.10 in Morycowa & Olszewska

2013) probably belong to the genus Cornuspira because they

also do not possess the diagnostic trait of Spirillina, i.e., glo-

bular proloculus followed by a gradually enlarging enrolled

undivided tubular second chamber. The specimen determined

as ?Rumanolina sp. shown in fig. 4.8 probably belongs to

Trocholina (T. ultraspirata Blau, 1987a) that occurs in the Lower

Jurassic (Rigaud et al. 2013).

After our revision, it seems that several species of forami-

nifers of the Vršatec Limestone appeared in the Tethyan Realm

for the first time during the Bajocian (Hungarillina lokutiense,

Radiospirillina umbonata, Ophthalmidium caucasicum,

O. ter quemi, O. obscurum, Paalzowella turbinella, Cornuspira

tubicomprimata, Nubecularia reicheli), or do not stratigraphi-

cally extend into the Bathonian (Tethysiella pilleri). Although

communities with benthic foraminifers with calcareous tests

(porcelaneous high-Mg calcite miliolids and mono- or poly-

crystalline low-Mg calcite spirillinids) are usually not used as

stratigraphic markers in the Jurassic successions, their compo-

sition and co-occurrence patterns allow an accurate dating of

the Vršatec Limestone. Therefore, in contrast to Morycowa &

Olszewska (2013), we argue that assemblages with benthic

foraminifers rather indicate that biohermal and peribiohermal

limestones of the Vršatec Limestone developed during

the Bajocian. This stratigraphic inference is in accordance

with stratigraphic data on ammonites that occur in dykes

within the Vršatec Limestone at Vršatec-Castle (Nanno lyto

ceras tripartitum) (Schlögl et al. 2009a). The co-occurrence of

Hungarillina lokutiense (first occurrence (FO) Bajocian),

Radiospirillina umbonata (FO Bajocian), Ophthalmidium

caucasicum (FO Bajocian), Ophthalmidium terquemi (FO

Bajocian), Paalzowella turbinella (FO Bajocian), Cornuspira

tubicomprimata (FO Upper Bajocian), Ophthalmidium

obscurum (FO Upper Bajocian), and Tethysiella pilleri (last

occurrence (LO) Bajocian) fully substantiates their Bajocian

age (Fig. 9).

The comparison between the lower and upper parts of

the Vršatec Limestone at Vršatec-Javorníky indicates that

the overall composition remained similar, being dominated by

Nubecularia and Ophthalmidium (Fig. 10), and no major

stratigraphic turnover is thus apparent within the Vršatec

Limestone. Dykes of crinoidal limestones penetrate into coral

limestones and bedded crinoidal limestones directly overlie

coral limestones at Vršatec-Javorníky. At Vršatec-Castle,

the thickness of crinoidal limestones is locally extremely thin,

reduced to 2.5 m or even zero, and the Bohunice Formation

locally directly overlies peribiohermal facies (with breccias)

on the top of the Vršatec Limestone. The upper boundary of

the Vršatec Limestone is pre-dating the termination of cri-

noidal limestones at some sites or temporally coincides with

the termination of crinoidal limestones of the Krupianka and

Smolegowa formations. Thus the deposition of the Vršatec

Limestone clearly terminated prior to the middle Garantiana

Zone (Late Bajocian) as suggested by maximum strati-

graphical extent of the crinoidal limestones on the Czorsztyn

Ridge. However, the termination of coral reefs probably

occurred earlier. First, the boundary between the coral lime-

stones and crinoidal limestones is consistently characterized

by a hard ground surface, indicating a major hiatus in depo-

sition. Second, the presence of several tens of meters of

crinoidal limestones (which consistently terminated their

deposition during the early Garantiana Zone in the Pieniny

Klippen Belt, Wierzbowski et al. 2004) that overlie coral

limestones at Vršatec-Javorníky indicate that the coral-reef

production stopped prior to the Garantiana Zone. We note

that for mation of coral reefs largely terminated at the end

of the Humphriesianum zone in western Europe (Durlet et al.

2001).

Paleogeographic comparison of foraminifer communities

The foraminifer assemblages of the Vršatec Limestone are

dominated by calcareous miliolid, spirillinid, and involutinid

species. Differences in the composition of assemblages

between the two Vršatec sites (with dominance of miliolids)

on one hand and Malé Hradište on the other hand (dominated

by Paalzowella) indicate some degree of environmental hete-

rogeneity among sites (Fig. 11). Similar types of assemblages

Fig. 9. Stratigraphic ranges of foraminifer species described in this

study overlap in the Bajocian stage. The upper stratigraphical extent

of K. cf. altissima and T. turris are based on the Bajocian age of

the Vršatec Limestone as documented in this paper.

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Assemblages dominated by encrusting miliolids

(Nubecularia, Vinelloidea), mobile miliolids (Ophthalmidium),

and spirillinids (Paalzowella, Tethysiella, Radiospirillina,

Hungarillina) characterized carbonate platforms with corals

from the French Jura and Burgundy (and Subalpine Basin)

(Piuz 2004; Clerc 2005). Shallow-platform assemblages in

coral habitats were dominated by spirillinids, Paalzowella,

Tethysiella, low-energy deep-platform assemblages were

dominated by miliolids (Labalina, Ophthalmidium,

Cornuspira), nodosariids, and by agglutinated foraminifers,

and high-energy crinoidal and shelly assemblages were domi-

nated by Nubecularia and Lenticulina. Some miliolids such as

Ophthalmidium have a broad environmental distribution not

limited to a single substrate or depth. It inhabited coral-free

pelagic carbonate platforms (Böhm et al. 1999) as well as

coral and crinoidal environments on carbonate platforms (Piuz

2004). Some species of Nubecularia participate in the cortex

of oncoids on shallow carbonate platforms (Rat 1966), some

others formed large oncoids in aphotic environments of

pelagic carbonate platforms (Gradziński et al. 2004), and con-

sortia with microbes in deep-shelf environments with sponges

(Reolid 2011). However, diversity of foraminifer assemblages

in these oncoidal assemblages tends to be smaller than diver-

sity of assemblages from the Vršatec Limestone. The co-occur-

rence of different morphogroups and the relatively high genus

diversity show close similarity to those foraminifer assembla-

ges inhabiting carbonate platforms of Burgundy and French

Jura during the Bajocian.

Conclusions

The research integrating field data, micro- and macro-

paleontological taxonomic analyses and paleoecological

ana lyses of coral bioherms and peri-biohermal deposits of

the Vrša tec Limestone shows significant evidence for their

Bajocian age (proposed previously on the base of ammonites

from the dykes infillings and brachiopod and coral associa-

tions), in contrast to previous studies suggesting an Oxfordian

age. In this light, the Vršatec klippen consist of one single,

continuous Middle Jurassic–Lower Cretaceous succession,

rather than representing two tectonic slices with different

depositional histories. The benthic foraminifer assemblages of

coral limestones contain several species that have either their

first or last appearance during the Bajocian. This study thus

shows that analyses based on the distribution of benthic fora-

minifers in thin-sections represent a powerful tool for

the biostratigraphic dating, especially if biostratigraphically

important groups such as ammonites are absent. The composi-

tion and diversity of the benthic foraminifers of the Vršatec

Limestone is similar to the Bajocian carbonate-platform envi-

ronments with corals of the French Jura and Burgundy.

Acknowledgements: We thank to Felix Schlagintweit and

an anonymous reviewer for critical comments and Matúš

Hyžný for editorial comments. This study was financially

supported by the Slovak Academic Information Agency

(SAIA), the Slovak Scientific Grant Agency (VEGA 0169-19),

and the Slovak Research and Development Agency (APVV 17-

0555).

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