www.geologicacarpathica.sk
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
, OCTOBER 2011, 62, 5, 413—433 doi: 10.2478/v10096-011-0030-y
Introduction
Pelagic Jurassic to Cretaceous sediments are exposed close
to the village of Lókút, Bakony Mts, Hungary. These rocks
are rich in invertebrate macrofossils which were originally
collected by a team from the Geological Institute of Hungary,
in the early 1960s. The collection work was supervised by
the late Prof. József Fülöp. Among the macrofossils of the
Lókút profile belemnites were collected, but have remained
unstudied until now. A biostratigraphical framework is pro-
vided for the Tithonian by ammonites (Vigh 1984) which is
revised and updated in this paper. More recently Grabowski
et al. (2010) provided a stratigraphical scheme based on
magneto- and calpionellid-stratigraphy.
The aim of this study is: (a) to provide an accurate and up-
dated biostratigraphy for the ammonite rich Upper Jurassic
(?Oxfordian—Tithonian) part of the Lókút section, (b) to doc-
ument the diverse belemnite fauna, as no belemnites were
previously described from this stratigraphic interval from
Hungary, and (c) to perform oxygen and carbon isotope
analyses on belemnite calcite to compare with the biostrati-
graphical data and with previously published isotope curves.
Belemnites were studied and described by Nico M.M.
Janssen, István Főzy summarized ammonite biostratigraphy,
High-resolution ammonite, belemnite and stable isotope
record from the most complete Upper Jurassic section of the
Bakony Mts (Transdanubian Range, Hungary)
ISTVÁN FŐZY
1
, NICO M.M. JANSSEN
2
and GREGORY D. PRICE
3
1
Department of Paleontology and Geology, Hungarian Natural History Museum, P.O. Box 137, H-1431 Budapest, Hungary;
fozy@nhmus.hu; semiformiceras@gmail.com
2
Geertekerkhof 14bis, 3511 XC Utrecht, The Netherlands; hibolithes@hotmail.com
3
School of Geography, Earth & Environmental Sciences, University of Plymouth, Drake Circus, Plymouth, PL4 8AA, United Kingdom;
g.price@plymouth.ac.uk
(Manuscript received November 9, 2010; accepted in revised form March 17, 2011)
Abstract: This research focuses on the cephalopod fauna and biostratigraphy of the latest Jurassic succession of the
Lókút Hill (Bakony Mts, Transdanubia, Hungary). Fossils were collected bed-by-bed from Ammonitico Rosso facies
and from the subsequent Biancone type rock. The poorly preserved cephalopods from the lowermost part of the profile,
immediately above the radiolarite, may represent a part of the Oxfordian stage. The rich Kimmeridgian ammonite fauna
is published for the first time while the formerly illustrated Tithonian fauna is revised. All the successive Kimmeridgian
and Early Tithonian Mediterranean ammonite zones can be traced. The highest documented ammonite zone is the Late
Tithonian Microcanthum Zone. The beds above yielded no cephalopods. Particular attention was paid to the belemnite
fauna of over 120 specimens collected under strict ammonite control. Among the belemnite faunas an Early Tithonian,
an early middle Tithonian, a late middle Tithonian, and a latest Tithonian assemblage can be distinguished. Thereby, an
association is distinguished in the middle Late Kimmeridgian and one that characterizes the Oxfordian-Kimmeridgian
boundary beds. The main difference from previously published belemnite data appears to be that the Hungarian assem-
blages are impoverished with respect to contemporary faunas from Italy and Spain (Mediterranean Province). An isoto-
pic analysis of the belemnites show that the carbon-isotope data are consistent with carbon-isotope stratigraphies of the
Western Tethys and show a decrease in values towards the Jurassic-Cretaceous boundary.
Key words: Late Jurassic, Hungary, Bakony Mts, biostratigraphy, Ammonitico Rosso, ammonites, belemnites, stable
isotopes.
while isotope analyses and interpretations were undertaken
by Gregory D. Price.
Geological setting
The studied section is situated in the south-western part of
the central Bakony Mts, about 600 meters from the village of
Lókút, on the south-western edge of the Lókút Hill (Fig. 1).
The outcrop is in the Transdanubian Range, representing a
part of the Bakony Unit in the Austroalpine part of the
AlCAPA (Alpine-Carpathian-Pannonian) composite terrain
(Csontos & Vörös 2004). Lókút Hill is an exceptional place,
where 7 of the 11 Jurassic stages are represented by means of
macrofossils (mainly ammonites). The succession of the
Lókút Hill is the most complete and thickest Jurassic succes-
sion of Transdanubia, deposited in a deep, pelagic environ-
ment, within a “horst and graben” context (Galácz & Vörös
1972). The Hettangian-Bajocian rocks are exposed in a long
artificial trench and in a pit on the southern part of the hill
(Galácz 1975; Géczy 1976). The Upper Jurassic succession,
which is in the focus of our study, is exposed in a shallow,
20 meter long artificial trench on the southern slope of the
Lókút Hill. The geographical coordinates for the section are
414
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
47°12
’17”N, 17°52’56”E. Strata dip to north with mean
orientation 360°/20°.
In the lowermost part of the section the cherty Lókút Radi-
olarite Formation crops out. This is succeeded by 0.1 m of
light red-brown clay, followed by a light red-brown cherty
limestone. The following few meters consist of red and yel-
lowish, thick-bedded nodular limestone (the Pálihálás Lime-
stone Formation), which passes continuously into light grey
coloured, less nodular ammonite rich facies (the Szentiván-
hegy Limestone Formation). The uppermost part of the pro-
file is formed by the white, thin-bedded, Biancone type
limestone (of the Mogyorósdomb Limestone Formation),
which continuously develops from the underlying formation.
No sharp boundaries between the three carbonate formations
are apparent. A brief description of the above-mentioned
lithostratigraphical units are given in Császár (1997).
Stratigraphy
The cephalopods were gathered in 1962—1964. Because
the original field notes are lost, the only available informa-
tion on the sampling are the bed numbers written on the la-
bels for each fossil and the thickness data of the beds in Vigh
(1984). Vigh published the Tithonian ammonites of the sec-
tion and also noted the presence of the Kimmeridgian strata.
No details of the Kimmeridgian were, however, provided by
Vigh (1984). Although it is not known exactly where the
Bed 1 was placed in the section by Vigh (1984) as no obvi-
ous marker bed exists to anchor the succession, we were able
to reconstruct the bed numbering after re-examination of the
succession and comparing thickness, lithology and fossil
content. Furthermore, we have recently resampled a number
of horizons of the section to gather further ammonite data.
Grabowski et al. (2010), recently published magneto- and
biostratigraphical schemes of the upper part of the section.
They were able to recognize magnetochrons M21r through
to 18r spanning the Jurassic-Cretaceous boundary. They also
undertook calpionellid studies and on this basis they as-
signed the uppermost 4.5 meters of the section to the Berria-
sian. Consequently the studied part of the Lókút profile
represents a succession spanning the ?Late Oxfordian—Kim-
meridgian to earliest Berriasian.
Cephalopod fauna
Ammonites were preserved as only internal moulds
throughout the section. As the fauna consists of solely Medi-
terranean (Tethyan) elements, the zonation scheme of Enay
& Geyssant (1975) and Olóriz (1978) was used. Vigh (1984)
described only the Tithonian part of the ammonite succes-
sion. In this paper we not only re-evaluate his data, but we
also give details on the possible Late Oxfordian and on the
Kimmeridgian fauna as well. The stratigraphic distribution
of the most characteristic ammonites is given in Table 1.
The belemnite material studied is of varying quality. Many
belemnites appear to be incomplete or fragmentary, probably
as a result of the rigidity of the host rocks. In the reddish co-
loured ?Oxfordian—Kimmeridgian part (Beds 68 to 77) belem-
nites appear to be partially corroded. The stratigraphic
distribution of the belemnites is given in Table 2.
The Tithonian cephalopods from the Lókút section are de-
posited in the Museum of the Hungarian Geological Insti-
Fig. 1. The geographic (A) and tectonic position of the Lókút section (B). The location of the studied profile is marked by an asterisk. Map
‘B’ is modified Haas et al. (2006) showing the structural units of the pre-Cenozoic basement of the Pannonian Basin and the surrounding
mountain ranges.
415
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Table 1: Distribution of the diagnostic Late Jurassic ammonites in the Lókút section. Phylloceratids and lytoceratids are not indicated.
Haploceras elimatum and related microconch forms (H. carachtheis and associated species), although common throughout the Tithonian,
are also not shown. (Haplocer. – Haploceratidae; H. – Himalayitidae.)
416
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
tute, while the Kimmeridgian fossils were deposited to the
collection of the Paleontological Department of the Hungarian
Natural History Museum.
?Oxfordian
The lowermost two beds above the radiolarite (Beds 76, 77)
yielded no ammonites only belemnites. It may indicate that
these beds were deposited between the aragonite compensa-
tion depth and carbonate compensation depth. Some of these
belemnites (Subulibelus) were previously described from the
Oxfordian-Kimmeridgian levels of southwest Germany
(Riegraf 1981). Others, like Hibolithes semisulcatus, are
known to first occur in the latest Oxfordian (Riegraf 1981),
probably after the Epipeltoceras bimammatum (Sub)Zone.
Since the Oxfordian is generally represented by the cherty
formation of the Lókút Radiolarite, Oxfordian ammonites are
rare in the Transdanubian Range. They are missing from other
Bakony localities, and can be found only in the eastern part of
the range, in the Gerecse Mts (Főzy & Meléndez 1997).
Kimmeridgian
Upwards in the sequence, from Bed 75, the following ca. 3
meters of reddish, clayey, ammonite rich marls are Kimmeridg-
ian in age. All of the Kimmeridgian Mediterranean ammonite
zones can be recognized. The stratigraphy is based on more
then 200 ammonite specimens (Table 1), although many of
them are preserved only as fragments. The lower part of the
stage is rather condensed, hence the individual zones are repre-
sented by individual beds. Since the Kimmeridgian ammonite
successions are poorly known all over the Mediterranean Pro-
vince, and the usage of the ammonite zones is not obvious, the
zonal boundaries in the Lókút section are also tentative. Among
phylloceratids, Sowerbyceras is the most abundant, principally
in the Lower Kimmeridgian beds. Oppeliids, especially tara-
melliceratids are diverse. Taramelliceras strombecki and T.
compsum, index forms of the Strombecki and Compsum Zones
are represented by some well preserved specimens. Nebrodites
and related genera (Lessiniceras, Trenerites) are very common
especially in the lower and middle part of the Kimmeridgian
strata. The extremely evolutely coiled Nebrodites cavouri is
represented by a single, but well preserved specimen. Aspi-
doceratids are also abundant but because their taxonomy is still
problematic, their stratigraphic value is also rather limited. It
seems that the flat sided, moderately evolute Aspidoceras wolfi,
and the very globulose Physodoceras avellanum characterize
the earliest Kimmeridgian interval. The common and moderate-
ly inflated, variably forms with a single or two rows of tubercles
(Orthasidoceras and Aspidoceras s.str.) were not studied in de-
tail. The evolute and flat shaped genera with or without ventral
groove (Hybonoticeras and Pseudowaagenia) bloom in the up-
permost Kimmeridgian Beckeri Zone. The zonal index was not
found, but H. pressulum, which has a very similar range, is
present. Among perisphinctids, the presence of the rarely re-
corded Early Kimmeridgian Pseudosimoceras stenonis and the
abundance of the also poorly known Trapanasites adelus is
noteworthy. Some of the important Kimmeridgian ammonites
are illustrated in Figs. 2—4.
Table 2: Distribution of the Late Jurassic belemnites from Lókút
section.
417
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Fig. 2. Representative Kimmeridgian ammonites from the Lókút section. 1, 2 – Physodoceras wolfi (Neumayr, 1873) (Inventory number:
M 92 852) Bed 75; 3, 4 – Taramelliceras strombecki (Oppel, 1858) (M 92 879) Bed 73; 5, 6 – Hemihaploceras schwageri (Neumayr,
1873) (M 82 931) Bed 68; 7, 8 – Trapanesites adelus (Gemmellaro, 1872) (M 92 886) Bed 68.
418
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Fig. 3. Representative Kimmeridgian ammonites from the Lókút section. 1, 2 – Taramelliceras compsum (Oppel, 1863) (Inventory num-
ber: M 92 888) Bed 68; 3 – Streblites cf. tenuilobatus (Oppel, 1863) (M 92 1097) Bed 73; 4, 5 – Pseudowaagenia acanthomphala (Zittel,
1870) (M 92 1131) Bed 65; 6, 7 – Trenerites cf. evolutus (Gemmellaro, 1876) (M 92 958) Bed 73.
419
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Fig. 4. A representative Kimmeridgian ammonite from the Lókút
section. 1, 2 – Nebrodites cavouri (Gemmellaro, 1872) (M 92
1247) Bed 69.
Among the Kimmeridgian belemnites Mesohibolitidae are
most abundant but overall belemnites are rather rare, appar-
ently being restricted to the earliest Kimmeridgian and the
middle part of the Late Kimmeridgian. The middle Late
Kimmeridgian beds contain both Duvaliidae and Mesohibo-
litidae. As Late Kimmeridgian duvaliid belemnites are not
known yet, the fauna is of particular interest. Unfortunately,
the material is not well preserved and rather incomplete, ham-
pering any certain specific determination. The earliest Kim-
meridgian delivered a more abundant and diverse fauna. Here
Mesohibolitidae (Acutibelus, Hibolithes and Subulibelus?)
are clearly most abundant but some duvaliids do occur too
(Produvalia aff. nicosiai).
Tithonian
The upper 61 beds of the red, nodular limestone, which be-
come lighter coloured, nearly white up sequence, represent
the Tithonian stage. All of the Early Tithonian ammonite
zones were recognized, while the Late Tithonian is repre-
sented by the Microcathum Zone only. No younger cephalo-
pods were found in Lókút.
Among phylloceratids, Ptychophylloceras (especially P.
ptychoicum) is the most common. Lytoceratids, like the me-
dium sized Protetragonites are also abundant, however big-
ger forms (Lytoceras and it’s allies) are common too. With
respect to the Ammonitina, the genus Haploceras (especially
H. elimatum and associated forms) is very frequent. It makes
up 23 % of the whole Tithonian fauna (Vigh 1984). The
stratigraphy is based on more then 70 ammonite specimens,
which are indicated on Table 1.
The lower part of the Tithonian succession contains some
poorly preserved fragments of perisphinctids and aspidocer-
atids, including thickly ribbed Hybonoticeras, indicating the
base of the lowermost Tithonian Hybonotum Zone (Beds
58—61). The presence of Virgatosimoceras cf. albertinum in
Bed 61 is somewhat controversial. The specimen (figured by
Scherzinger et al. 2010: fig. 5.2) is thought to be characteristic
for the subsequent Darwini Zone ( = Albertinum Zone in
Olóriz 1978). This zone is represented by Beds 54—57, which
also yielded a rather poor fauna, mainly of perisphinctids and
aspidoceratids. The base of the zone was drawn by the appear-
ance of Haploceras cassiferum. This ammonite, which is of-
ten regarded as a macroconch of H. verruciferum, (Cecca &
Enay 1991), the characteristic ammonite of the subsequent
ammonite zone (Semiforme / Verruciferum), is however often
found, below the Semiforme Zone. The middle Tithonian
Semiforme, Fallauxi and Ponti Zones were documented on the
basis of diagnostic haploceratids, aspidoceratids, and simocer-
atids. The base of the Semiforme Zone (Beds 45—53) is
marked by the simultaneous appearance of H. verruciferum
and Volanoceras aesinense. The zonal index was found only
in debris. Densely ribbed Discosphinctoides rhodaniforme,
Pseudohimalayites and Virgatosimoceras are also present and
characteristic. The latter is represented by a recently described
species (V. dunaii) which fills the gap between the earlier
known older and younger chronospecies (Scherzinger et al.
2010). The subsequent Fallauxi Zone (Beds 27—44) contains a
diverse simoceratid fauna which clearly indicate the presence
of the upper part of the zone (Biruncinatum/Admirandum
Subzone). Simoceras admirandum and allied forms are rela-
tively frequent, while typical Simoceras biruncinatum was not
found in Vigh’s collection. It was gathered only during our re-
cent collecting work. The lower part (Richteri Subzone) can-
not be documented by means of ammonites. Fragments of the
large sized simoceratid (Volanoceras perarmatiforme) already
mark the Ponti Zone (Beds 24, 25). Beds above yielded a
moderately diverse fauna, including rare Simospiticeras and
some specimens of Moravisphinctes and Paraulacosphinctes.
Some of the stratigraphically important Tithonian ammonites
are illustrated on Figs. 5, 6.
On the basis of the distribution of the belemnites the Ti-
thonian sediments can be divided into four belemnite assem-
blages. The base of the Tithonian is characterized by
“Pseudobelus” zeuschneri while Hibolithes semisulcatus is
common. This low relative diversity assemblage is called
TiBA-I and encompasses the Beds 55 to 61. According to the
accompanying ammonites this assemblage corresponds to the
Hybonoticeras hybonotum and the Semiformiceras darwini
Zones. The next belemnite assemblage (TiBA-II; Beds 41 to
46) yields Duvalia ensifer and Conobelus strangulatus. It is
found in the top of the Semiformiceras semiforme Zone and
the base of the Semiformiceras fallauxi Zone. Duvalia cf.
abeli and common H. semisulcatus further characterize this
assemblage. “Pseudobelus” ex gr. zeuschneri and Hibolithes
conradi occur in the latest Semiformiceras fallauxi Zone
(Simoceras admirandum Subzone) to Micracanthoceras
microcanthum Zone. This part encompasses the Beds 16 to 32
and can be divided into two successive belemnite assemblages
(TiBA-III and IV; Table 2). TiBA-III furthermore yields
Hibolithes cf. fellabrunnensis, Duvalia cf. esba? and the last
Hibolithes semisulcatus. TiBA-IV yields Hibolithes conradi,
“Duvalia” tithonia and the last “Pb.” ex gr. zeuschneri.
420
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Fig. 5. Representative Tithonian ammonites from the Lókút section. 1 – Lemencia cf. pergrata (Schneid, 1914) (J 10.60.1) Bed 38; 2 – Pseudo-
lissoceras olorizi Főzy, 1988 (J 9769) Bed 54; 3, 4 – Paraulacosphinctes sp. (J 10.61.1) Bed 20; 5 – Neochetoceras sp. (J 10879) Bed 43.
6 – Haploceras cassiferum Főzy, 1988 (J 10.63.1) Bed 57; 7 – Subplanitoides sp. (J 10.62.1) Bed 44; 8 – Simospiticeras lojense Olóriz
& Tavera, 1977, (J 10.64.1) Bed 10; 9, 10 – Discosphinctoides rhodaniforme Olóriz, 1978 (J 10.65.1) Bed 53.
421
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Fig. 6. Representative Tithonian ammonites from the Lókút section. 1, 2 – Semiformiceras semiforme (Oppel, 1865) (Inventory no:
J 10.54.1) from loose; 3 – Volanoceras perarmatiforme (Schrauroth, 1865) (J 10.55.1) Bed 26; 4 – Volanoceras aesinense (Meneghini,
1885) (J 9778) Bed 53; 5 – Simolytoceras sp. (J 10.56.1) Bed 24; 6, 7 – Orthaspidoceras sp. (J 10.57.1) Bed 54; 8, 9 – Physodoceras
neoburgense (Oppel, 1863) (J 10.58.1) Bed 55; 10, 11 – Hybonoticeras hybonotum (Oppel, 1863) (J 10.59.1) Bed 59.
422
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Three of the four assemblages are separated by beds that
apparently contain virtually no belemnites, the Beds 47 to 54
and the Beds 33 to 40. So do the Beds 62 to 67 that separate
the first Tithonian assemblage from the very condensed beds
that contain ?Oxfordian—Kimmeridgian cephalopods. Some of
the Tithonian belemnites from Lókút are illustrated in Fig. 7.
Stable isotopes
Methodology
Stable isotopes were determined on a VG Instruments Opti-
ma Isotope Ratio Mass Spectrometer with a Multiprep Auto-
mated Carbonate System (at the University of Plymouth) using
200 to 300 micrograms of carbonate. Isotopic results were cali-
brated against NBS-19. Reproducibility for both
18
O and
13
C
was better than ± 0.1 ‰, based upon multiple sample analysis.
All belemnite samples were analysed additionally for trace ele-
ment contents to evaluate diagenetic alteration. Thin section
analysis in conjunction with trace element analysis was used to
determine the state of preservation of each of the fossil types ex-
amined. Prior to chemical and isotopic analysis, areas of each
belemnite deemed most susceptible to diagenetic alteration,
principally the exterior of each shell were removed. The re-
mains were fragmented, washed in ultra pure water and dried in
a clean environment. Fragments were subsequently picked un-
der a binocular microscope to secure those judged to be best
Fig. 7. Representative Tithonian belemnites from the Lókút section. 1, 2 – Hibolithes ex gr. semisulcatus (Münster, 1830) (Inventory no:
J 10.1.1) Bed 46; 3, 4 – Conobelus strangulatus (Oppel, 1865) (J 10.2.1) Bed 41; 5, 6 – “Pseudobelus” ex gr. zeuschneri (Oppel, 1865)
(J 10.3.1) Bed 32; 7 – Hibolithes cf. fellabrunnensis (Vetters, 1905) (J 10.4.1) Bed 32; 8, 9 – “Pseudobelus” zeuschneri (Oppel, 1865) juv.
(J 10.5.1) Bed 61; 10, 11 – Hibolithes conradi Kilian, 1889 (J 10.6.1) Bed 24; 12, 13 – Duvalia cf. abeli (Vetters, 1905) (J 10.7.1)
Bed 46; 14, 15 – Duvalia ensifer (Oppel, 1865) (J 10.8.1) Bed 45; 16, 17 – “Pseudobelus” zeuschneri (Oppel, 1865) (J 10.5.2) Bed 61;
18, 19 – Hibolithes semisulcatus (Münster, 1830) (J 10.9.1) Bed 25; 20, 21 – Duvalia cf. esba (de Gregorio, 1885) (J 10.10.1) Bed 32;
22 – Hibolithes ex gr. semisulcatus (Münster, 1830) (J 10.11.1) Bed 45; 23 – Hibolithes ex gr. semisulcatus (Münster, 1830) (J 12.12.1)
Bed 58; 24, 25 – “Pseudobelus” zeuschneri (Oppel, 1865) (J 10.13.1) Bed 58.
423
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Table 3: Isotopic and elemental compositions of belemnite rostra from Lókút.
Belemnite species
Bed
number
δ
13
C
(PDB)
δ
18
O
(PDB)
Ca
(ppm)
Fe
(ppm)
Mg
(ppm)
Mn
(ppm)
Sr
(ppm)
Hibolithes cf. conradi*
Bed 16
0.9
1.4
371684
100
1890
64
240
Duvalia tithonia
Bed 18
–1.7
–1.5
355032
60
2026
29
1049
H. cf. conradi
Bed 18
–1.8
–0.9
359047
109
2317
20
972
Pseudobelus ex.gr. zeuschneri Bed 21
–1.0
–0.2
350100
17
1799
5
895
Belemnites sp.*
Bed 21
–0.3
–1.8
391288
68
1384
33
234
H. cf. conradi*
Bed 22
1.4
–0.4
356897
23
1615
26
270
Hibolithes sp.*
Bed 23
0.6
–1.2
385583
195
1963
179
208
H. cf. conradi
Bed 24
–1.6
–1.3
351484
26
2567
11
1055
Hibolithes sp. *
Bed 24
1.2
–2.6
361603
67
1993
140
525
Pseudobelus ex.gr. zeuschneri Bed 25
–2.3
–2.9
353975
44
2151
24
898
H. cf. semisulcatus*
Bed 25
–1.9
–2.2
349514
23
2231
104
794
H. cf. semisulcatus*
Bed 30
–0.8
0.4
359052
53
2758
125
1002
Pseudobelus ex.gr. zeuschneri Bed 32
–1.2
–0.9
351008
18
1860
10
955
Hibolithes sp.
Bed 32
–0.8
–1.1
362398
29
2431
9
995
H. cf. fellabrunnesis
Bed 32
–0.6
–1.3
362530
32
2503
26
864
Duvalia cf. ensifer*
Bed 32
0.4
0.4
388057
87
1939
591
470
H. cf. semisulcatus
Bed 35
–1.4
–1.6
361858
22
2525
18
1119
Hibolithes semisulcatus*
Bed 40
–1.2
–2.4
361213
72
1601
368
484
Conobelus cf. strangulatus*
Bed 42
–1.0
–1.4
348063
293
1936
435
482
H. cf. semisulcatus*
Bed 42
0.8
–1.2
349304
73
1933
69
536
Hibolithes sp.
Bed 43
–1.8
–2.2
365793
41
1861
20
1020
Duvalia cf. ensifer
Bed 44
–0.2
–0.1
358767
93
2004
14
908
Conobelus cf. strangulatus*
Bed 44
–0.8
–1.7
344693
121
1937
111
860
Hibolithes cf. semisulcatus*
Bed 44
–0.4
–1.1
328534
136
2240
248
801
Duvalia ensifer*
Bed 45
–3.2
–2.1
365216
82
1365
115
334
Conobelus cf. strangulatus*
Bed 45
0.0
–1.0
358167
61
2255
734
690
Duvalia ensifer*
Bed 45
1.3
–0.8
363919
493
2585
134
442
Hibolithes sp. *
Bed 45
–0.8
–0.9
355709
19
2253
99
939
Hibolithes semisulcatus*
Bed 45
1.3
–0.7
331695
152
2015
55
680
Hibolithes semisulcatus
Bed 46
–0.7
–1.1
339289
14
2286
11
771
Conobelus cf. strangulatus
Bed 46
–1.5
–1.2
351746
31
2166
19
1016
H. cf. semisulcatus
Bed 46
–0.9
–1.2
329436
34
2030
42
955
Conobelus cf. strangulatus*
Bed 46
–1.5
–0.1
343629
61
1792
68
996
Duvalia cf. abeli*
Bed 46
1.9
–1.2
347496
318
2163
118
215
Duvalia cf. ensifer*
Bed 46
–1.2
–0.4
362948
24
1738
84
981
Duvalia cf. ensifer*
Bed 46
–0.5
0.3
361830
24
2171
84
1029
Hibolithes sp.*
Bed 47
–0.1
–1.3
356315
195
1834
869
458
Pseudobelus cf. zeuschneri
Bed 55
0.4
–1.2
356031
93
1827
8
989
H. cf. semisulcatus*
Bed 55
0.3
–1.6
363467
55
2851
68
1097
Hibolithes sp.
Bed 56
1.8
–1.4
381341
17
2627
36
932
Hibolithes sp.*
Bed 57
0.3
–0.5
346926
128
1927
684
713
Pseudobelus cf. zeuschneri
Bed 58
0.5
1.1
368994
76
1810
52
856
Hibolithes semisulcatus
Bed 58
1.4
–0.6
339196
80
1804
31
436
Hibolithes semisulcatus*
Bed 58
–0.4
–1.4
377196
69
2788
82
1131
Pseudobelus zeuschneri*
Bed 61
–0.7
–1.8
347528
63
1960
105
1067
Hibolithes sp. *
Bed 62
–0.1
–0.4
368284
75
2635
93
1077
Belemnites sp.
Bed 76
–0.3
0.5
365134
8
2328
8
855
Belemnites sp.
Bed 76
–0.6
0.2
369019
64
1626
8
918
Belemnites sp.
Bed 77
–0.5
0.3
370522
7
1500
6
901
* Deemed diagenetically altered
preserved, which were then analysed for oxygen and carbon
isotopes. Subsamples for chemical analysis weighing 5—10 mg
were dissolved in nitric acid and analysed using a Varian
725-ES Inductively Coupled Plasma spectrometer. According
to analysis of duplicate samples, reproducibility was better than
± 4 % of the measured concentration of each element.
Fossil preservation
The belemnites were not only fragmentary and frequently
displayed solution marks, as noted above, they were also often
‘chalky’ in appearance. After the
death of an organism, microbial
agents may attack the shells, di-
gesting the organic matrix that sur-
rounds the calcium carbonate
crystallites (Glover & Kidwell
1993; Brand 1994). The disassoci-
ation of these crystallites due to the
loss of the organic matrix holding
them together has an effect similar
to chemical dissolution and causes
pitting of the surface, chalkiness,
and loss of colour and/or lustre
(Glover & Kidwell 1993; Brand
1994). A chalky appearance alone
is however, not a reliable criterion
for defining the degree of diagenet-
ic alteration. Diagenetic alteration
was also apparent using standard
thin section analysis. Many rostra
examined (Fig. 8b—e) displayed a
mottled texture and possible
cloudy opaque dissolution resi-
dues. Mottling and cloudiness was
often concentrated around the api-
cal line as well the margins of the
rostra. Within those samples show-
ing a better state of preservation
(Fig. 8a,f) the calcite is generally
clear and primary growth lines are
apparent. Those rostra appearing
cloudy in hand specimen were also
those that displayed the mottling
and cloudiness seen in thin section.
Our trace elemental analysis is
consistent with these petrographic
observations in that they reveal the
poor state of preservation of many
of the belemnite rostra. Fe concen-
trations range from 15—493 ppm,
Mn
concentrations
from
5—
869 ppm, Mg concentrations from
1341—2851 ppm and Sr concentra-
tions from 186—1131 ppm in all 49
belemnite rostra analysed (Table 3).
Well-preserved belemnites typical-
ly show low concentrations of Mn
( < 50 ppm) and Fe ( < 200 ppm) and
higher concentrations of Sr (ca. 600—1600 ppm) (e.g. Nunn et
al. 2009; Price et al. 2009). Fe and Mn concentrations are typi-
cally higher in diagenetically altered calcite, as Fe
2+
and Mn
2+
are more soluble under reducing conditions and thus available
for replacing Ca
2+
in the calcite lattice (Brand & Veizer 1980;
Veizer 1983). Under such conditions there may also be a loss of
‘marine’ Mg and Sr (Veizer 1983). Hence many belemnites
were excluded on the basis of relatively high concentrations of
Mn and Fe or reduced concentrations of Sr and Mg potentially
reflecting dissolution as indicated by the chalky nature of many
of the belemnites.
424
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Isotope results
Apart from the isotope data from the lowermost (?Oxford-
ian) beds, the carbon isotope values of the belemnite calcite
through the Tithonian part of the section show a decline in
values from 1.8 ‰ in the Darwini Zone to —1.7 ‰ (V-PDB) in
the Microcanthum Zone. The belemnite oxygen isotopes show
some variability ranging from —2.9 to —0.1 ‰ (V-PDB). Obvi-
ous stratigraphic trends are not evident. Likewise although a
range of species were analysed (e.g. Hibolithes and Duvalia),
because of the generally poor state of preservation only two
Duvalia were deemed well-preserved. Hence potential eco-
logical trends were not discernable from the isotope data.
Fig. 8. Photomicrographs of belemnite rostra. a – Sample Belemnites sp., Bed 77 displaying clear and translucent growth lines. Scale bar is
1 mm. b – Sample Hibolithes semisulcatus, Bed 58 showing a mottled texture particularly concentrated around the apical line. Scale bar is 1 mm.
c – Sample Hibolithes sp., Bed 57 showing mottling and opaque cloudy texture particularly around the margin of the rostrum. Scale bar is 1 mm.
d – Sample Duvalia cf. abeli, Bed 46 shows a dense mottling and opaque cloudy texture. Scale bar is 1 mm. e – Sample Hibolithes cf.
semisulcatus, Bed 44 shows a mottled texture particularly concentrated around the apical line. Scale bar is 1 mm. f – Duvalia tithonia, Bed 18
shows generally clear calcite and primary growth lines with more opaque calcite around the margin of the rostrum. Scale bar is 0.5 mm.
The oxygen isotope data from the ?Oxfordian samples
show a narrow range (0.2—0.5 ‰ V-PDB). These data are a
little more positive than our Tithonian data.
Discussion
Ammonites
The Upper Jurassic ammonite fauna of the Lókút section
has a strong Mediterranean character, which is reflected by
the presence of numerous exclusively Tethyan ammonite
genera and species, as well as by the proportion of ammonite
425
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
which in general lack unambiguous traits. Hibolithes ex gr.
semisulcatus are well known from the Kimmeridgian-Titho-
nian boundary sediments and occur abundantly in both the
Submediterranean as in the Mediterranean Province, and oc-
casionally even in the Boreal-Atlantic Realm.
Duvalia-like belemnites are known from the Tethyan Late
Bathonian to Oxfordian, and probably the late Early Kim-
meridgian (Mariotti 2002a,b). Recently Hikuroa (2004) and
Challinor & Hikuroa (2007) published sedimentary succes-
sions from the Antarctic Peninsula which yielded Duvaliidae
from sediments of comparable ages. For the moment, there
appears to be a period (within the literature) from the Late
Kimmeridgian to earliest Tithonian in which the Duvaliidae-
do not occur. The “reappearance” of Duvaliidae (Conobelus
and duvaliid belemnites related to Duvalia ex gr. ensifer) co-
incides approximately with the boundary between the Lower
and middle Tithonian.
In comparing the Hungarian belemnite assemblages with
assemblages from Italy (Combémorel & Mariotti 1986a;
Mariotti 1995, 2002b, 2003), some discrepancies occur,
moreover the Hungarian assemblages appear to be impover-
ished. TiBA-IV contains belemnites that are characteristic
for the latest Jurassic. However, characteristic genera and
species like Conobelus and “Pb.” fischeri, are absent. Duvalia
tithonia is not know from older sediments than with calpionel-
lids that characterize the A2 Calpionellid Zone. Apparently,
our specimen occurs within the top of the Chitinoidella
Calpionellid Zone (cf. Grabowski et al. 2010).
Moreover, the Hungarian material indicate that much more
distinct assemblages emerge between the Early—middle Titho-
nian and the middle—Late Tithonian than previously indicated.
The assemblage described by Combémorel & Mariotti
(1986a) from the V. volanense Zone (Mariotti 1995) appears
to be comparable to TiBA-II in the Lókút section (top S. semi-
forme and base S. fallauxi Zones) but the Hungarian assem-
blage is clearly less diverse. In the Lókút section TiBA-I
encompasses almost the entire Early Tithonian (if distinguish-
ing between the Lower and middle Tithonian). The characteris-
tic belemnite (“Pb.” zeuschneri), would be the first indication
that duvaliids occur in the earliest Tithonian, shortening the pre-
viously mentioned interval of “lacking duvaliids”.
Duvaliids also occur in the Bed 68 (base Hybonoticeras
beckeri Zone). Due to the rigidity of the host-rock only the
transverse section can be studied, which shows an elongated
subquadrangular laterally flattened cross-section, typical for du-
valiid belemnites. Moreover, the Beds 68 and 70 yielded cono-
beloid belemnites, and in the Bed 70 one resembles Conobelus
massimoi (Mariotti, 2002). The latter has been described from
the Crussoliceras divisum Zone of northern Italy. In the
Beds 75 and 77 (probably Oxfordian-Kimmeridgian boundary
beds) duvaliids occur again.
Isotopes
Increasingly negative
18
O values in carbonates can be re-
lated to higher temperatures in environmental settings where
continental ice volume over time is relatively constant (and
therefore
18
O
seawater
is relatively invariable) and evaporation
or freshwater input are minor factors. Belemnites are also
suborders. As a result of this phylloceratids and lytoceratids
are very common throughout the section. For example, in the
Tithonian part of the succession these groups comprise about
60 % of the whole fauna (Vigh 1984).
Data for the Oxfordian are very scarce in Lókút, therefore it
cannot be discussed in detail. In contrast, the Kimmeridgian is
well developed and can be compared with some of the well-
documented Tethyan fauna, especially known from the Sub-
betics (Olóriz 1978), Southern Alps (Sarti 1984, 1993) and
Sicily (Pavia & Cresta, Eds. 2002). The Kimmeridgian suc-
cession of the Lókút Hill – although still condensed, as al-
ways in the Rosso Ammonitico facies – is the thickest and
most complete representation of the stage in Hungary, where
all Kimmeridgian ammonite zones can be traced.
Because the Tithonian in Lókút also shows a strong
Mediterranean affinity and its ammonite succession can be
compared easily with those known from other Tethyan locali-
ties, the novel zonal scheme, introduced by Vigh (1984) was
revised and replaced by a standard scheme. As in other locali-
ties of the Transdanubian Range the middle Tithonian is better
represented than the lowermost part of the stage. The Semi-
forme and Fallauxi Zones contain more beds then those of the
Hybonotum and Darwini Zones. The Ponti Zone is thin, but
the base of the Upper Tithonian (Microcanthum Zone) is char-
acterized by numerous beds and a relatively diverse and typi-
cal, but poorly known assemblage. This fauna contains some
little known ammonites, including flat, discoidal perisphinc-
tids with, or without a ventral groove (like Paraulacosphintes
and Oloriziceras), and some himalayitids (Microcanthoceras)
which are best known from the Subbetics (Tavera 1985).
Belemnites
The most recent information regarding Mediterranean Late
Jurassic belemnites originates from Italian sections (Combémo-
rel & Mariotti 1986a,b, 1990; Mariotti 1995, 2002a,b, 2003).
However, knowledge about the bed-by-bed distribution of
Tethyan belemnites is sparse (Mariotti 1995: p. 222—225).
Combémorel & Mariotti (1986a) described belemnites from the
Central Apennines. They dealt with belemnites from either the
S. semiforme Zone or the Volanoceras volanense Zone, although
later on Mariotti (1995) accepted a Volanense ( = Ponti) age for
the sediments. Cecca & Enay (1991) mentioned Duvalia ensifer
from the R. richteri Subzone, from Tithonian sediments in the
southeast of France. Combémorel & Mariotti (1986b) described
Duvalia tithonia from the A2 Calpionellid Zone and discuss to
some more extent stratigraphical details about this species.
Janssen (1997: text-fig. 2) described and mentioned some
belemnites from the latest Jurassic and earliest Cretaceous
from the southeast of Spain. However, especially the Conobe-
lus-species most probably are erroneous and additional mate-
rial (a.o. “Pseudobelus” fischeri Combémorel & Mariotti,
1990) still needs to be published. It appears, however, that
many belemnites that occur in TiBA-III and IV disappear in
the beds that straddle the Jurassic-Cretaceous boundary.
The latest Kimmeridgian to earliest Tithonian is character-
ized by belemnite faunas which are very low in diversity but
are generally very abundant (Riegraf 1981; Combémorel
1997). Most numerous are Hibolithes, a group of belemnites
426
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Fig. 9. Integrated stratigraphy of the Lókút section showing the correlation between biostratigraphic data and those based on magneto-
stratigraphy. The measured log is tentatively correlated with the bed numbers published by Vigh (1984). Correlation is based on known
thickness of the Tithonian part of the section, observed lithological changes and re-sampling of the profile. Ammonite zones for the Kim-
meridgian and Tithonian follow the standard zonation scheme by Enay & Geyssant (1975) and Olóriz (1978), respectively. Abbreviations:
(a) radiolarite (Lókút Fm), (b) red nodular limestone (Pálihálás Fm), (c) light coloured, thin-bedded limestone, with cherty layers (Szen-
tivánhegy Fm), (d) white, Biancone type limestone (Mogyorósdomb Fm). For the zonal names, from the oldest to the youngest: Platynota,
Strombecki, Divisum, Compsum Cavouri, Beckeri, Hybonotum, Darwini, Semiforme, Fallauxi, Ponti, Microcathum Zones. Belemnite As-
semblages are in accordance with the data published herein.
427
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
thought to have secreted their carbonate in isotopic equilibri-
um with ambient seawater (e.g. Lowenstam & Epstein 1954;
Price & Mutterlose 2004). This assumption is supported by
the fact that the
18
O values of modern Sepia shells (an extant
relative of belemnites) also appear close to equilibrium frac-
tionation (Bettencourt & Guerra 1999). Hence assuming oxy-
gen isotope equilibrium, relatively cool temperatures are seen
in the Oxfordian and warmer conditions in the Tithonian. The
18
O values of belemnites imply temperatures as low as 8 °C,
if a
seawater
value of —1.0 ‰ (SMOW), thought to be appropri-
ate for assumed non-glacial periods (Shackleton & Kennett
1975) using the paleotemperature equation of Anderson &
Arthur (1983) is used. According to the same assumptions the
warmest paleotemperatures reach 22 °C consistent with the
paleolatitude of the site. As the few Oxfordian samples were
derived from just above the Lókút Radiolarite, cooler condi-
tions may be indicative of a deeper depositional setting al-
though warmer conditions in the Tithonian are consistent with
data published from elsewhere (e.g. the Russian Platform,
Riboulleau et al. 1998; Price & Rogov 2009).
The carbon isotope data show a decrease through the Ti-
thonian part of the section that is consistent with carbon-iso-
tope stratigraphies of the Western Tethys (e.g. Weissert &
Channell 1989; Weissert & Mohr 1996) and also the Russian
Platform (Price & Rogov 2009) which show a decrease in
values towards the Jurassic-Cretaceous boundary. Weissert
& Channell (1989) interpret this
13
C decline as evidence of
increasingly oligotrophic conditions in the Tethyan seaway.
This pattern is therefore thought to reflect a global signal of
carbon cycling in the Late Jurassic oceans. Further stable
isotope data from the Lókút section are, however, required to
substantiate linkages between global records.
Conclusions
Re-measuring and re-sampling the section of the Lókút
Hill allowed us to allocate and anchor the cephalopod collec-
tion, gathered nearly 50 years ago.
Based on the rich and relatively well preserved, bed-by-bed
collected ammonite fauna, the biostratigraphical subdivision of
the lower, cephalopod bearing part of the Upper Jurassic—Lower
Cretaceous section was precisely done. Above the lowermost
(possibly Oxfordian) beds, a relatively complete succession of
the Kimmeridgian Platynota, Strombecki, Divisum, Compsum,
Cavouri and Beckeri Zones was recognized, which is followed
by the Tithonian Hybonotum, Darwini, Semiforme, Fallauxi,
Ponti and Microcanthum Zones.
The belemnite fauna – also collected under strict stratigraph-
ic control, together with the ammonites – allowed us to recog-
nize four belemnites assemblages (TiBA-I to TiBA-IV) for the
Tithonian part. We distinguished one assemblage in the middle
late Kimmeridgian and one for the ?Oxfordian-Kimmeridgian
boundary beds. However, due to the low amount and quality of
the material the latter two are only tentatively distinguished.
The Tithonian assemblages can be compared with previously
published belemnite assemblages but appear impoverished in
relation to Mediterranean (Italy, Spain) assemblages. The rather
species poor Kimmeridgian belemnite assemblage cannot be
compared because belemnite stratigraphical and taxonomical
details are missing from literature except for Mariotti (2002b,
2003). The latter author published a highly diverse assemblage
from the C. divisum Zone of northern central Italy.
The ?Oxfordian-Kimmeridgian assemblage which appears
rather diverse (mainly Mesohibolitidae) cannot be compared
in detail to other areas yet, because detailed stratigraphical
data are also missing. However, the general picture, repeated
in literature showing an abundance of Mesohibolitidae in the
Upper Oxfordian to Lower Tithonian sediments of the Medi-
terranean Tethys could actually be artificial due to lack in
well investigated sections, as demonstrated by Mariotti
(2002b, 2003). Moreover, we find indications that duvaliids
occur throughout the interval we investigated.
Despite the poor preservation of many of the rostra our
geochemical analyses of belemnite specimens from a strati-
graphically well-constrained section yielded new data for re-
constructions of paleoclimate and paleoecology. These data
point to a cool (or deeper) ?Oxfordian and a warmer Titho-
nian. Our
13
C data are consistent with other isotope stratig-
raphies and allude to the possibility of the Lókút section
recording global events.
Systematic section
On a family level there appear to be no large differences be-
tween latest Jurassic and Early Cretaceous belemnites, only
abundances fluctuate heavily. The belemnites described here
can be divided into two families, namely the Mesohibolitidae
and Duvaliidae. This division is based on the position of the
siphon towards the alveolar groove, being opposite in Duvali-
idae and situated on the same side in Mesohibolitidae. In addi-
tion the general morphology of the rostrum adds to this
separation, in which Duvaliidae are generally laterally flat-
tened, while Mesohibolitidae are rounded or dorso-ventrally
flattened and spindle-shaped (hastate or (sub)fusiform). How-
ever, the alveolar area of some Hibolithes can be laterally
compressed, as in Hibolithes conradi. Some suspected Du-
valiidae also show the same outer morphology during their
ontogenetical development, especially Conobelus. Thus ge-
neric attribution is not always definite, as the position of the
siphon is often not known. Moreover, the stratigraphical
record of the Duvaliidae is momentarily incomplete. There are
gaps in our knowledge regarding the phylogenetic relation be-
tween Late Jurassic and Early Cretaceous genera, although
morphological differences are often meagre. This is largely
the result of the near absence of published stratigraphical data
on belemnites from the Berriasian.
The main quandary is represented by the Tithonian belem-
nites attributed to the genus Pseudobelus. There is momen-
tarily no firm ground to place them in the genus created by
Blainville (1827) other than a morphological resemblance
(pers. observ. Janssen). For the moment they appear to belong
to a genus closely related to the Tithonian—earliest Cretaceous
duvaliids, or representing a genus between Produvalia and
Duvalia. These belemnites do show some characteristics of
Produvalia or Duvalia and of Pseudobelus. Yet, lateral lines
or incisions are often noted in other duvaliids too, and espe-
428
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
cially juvenile to immature specimens do tend to have the
same morphology as compared to pseudobeloid belemnites.
Pseudobelus s.s. are currently known to occur in uppermost
Berriasian to lower Upper Hauterivian sediments only.
Family: Mesohibolitidae Nerodenko, 1983
Genus: Acutibelus Riegraf, 1981
Acutibelus sp.
?1877 Belemnites semisulcatus Münster – Favre, pl. I, fig. 3a—b
M a t e r i a l : One incomplete specimen from Bed 75 and
one doubtful specimen from Bed 70.
D e s c r i p t i o n : A very elongated specimen with rather
faint alveolar groove. The juvenile or immature specimen
from Bed 70 shows a very deep and very sharp alveolus.
R a n g e : Early to earliest Late Kimmeridgian.
O c c u r r e n c e : Late Early to earliest Late Kimmeridgian
of southwest Germany (Riegraf 1981), Savoie (?Favre 1877)
and Hungary.
Genus: Hibolithes Denys de Montfort, 1808
Hibolithes conradi Kilian, 1889
(Fig. 7.10,11)
1868 Belemnites cfr. semisulcatus Münster – Zittel, pl. I, fig. 8
(fide Kilian, 1889)
?1871 Belemnites cfr. semisulcatus Münster – Gemmellaro, p. 21,
pl. III, figs. 2—3
1889 Belemnites (Hibolites) Conradi n. sp. Kilian, pp. 635, 690
(cf.), pl. XXVI, fig. 4
?1922 Belemnites Conradi Kilian – de Gregorio, p. 8, pl.I, fig. 12
pars 1990 Hibolithes semisulcatus (Münster): Combémorel & Mariotti,
pl. II, fig. 9
pars 1997 Hibolithes semisulcatus (von Münster) – Janssen, pp. 12—13
M a t e r i a l : Six incomplete specimens from Beds 24 to 16.
D e s c r i p t i o n : A robust rounded semihastate rostrum with
pointed, centrally placed apex. The alveolar area is character-
ized by lateral compression. Alveolus shallow with well de-
veloped wide alveolar groove. The apical line is central.
R a n g e : Latest Tithonian (M. microcanthum Zone) to
Late (?)Berriasian (Zone unknown).
O c c u r r e n c e : Hungary, Italy, Spain and (?)northern Alps.
Hibolithes cf. fellabrunnensis (Vetters, 1905)
(Fig. 7.7)
1905 Belemnites Fellabrunnensis Vetters, p. 245, text-fig. 1
M a t e r i a l : One incomplete rostrum from Bed 32 (the al-
veolar part is lacking).
D e s c r i p t i o n : Elongated rounded to slightly dorso-ven-
trally compressed rostrum with a relative long alveolar
groove, well on to the rostrum sollidum, and a shallow alve-
olus. The alveolar area is rounded (cf. Vetters 1905) but not
preserved in our material.
R a n g e : Late middle Tithonian (top S. fallauxi).
O c c u r r e n c e : Hungary and northern Alps.
Hibolithes semisulcatus (Münster, 1830)
(Fig. 7.1,2,18,19,22,23)
1830 Belemnites semisulcatus Münster, pp. 6—7, pl. 1, figs. 1—8, 15
1862 Belemnites diceratiana Etallon, p. 69
1870 Belemnites cfr. semisulcatus Münster – Zittel, p. 30, pl. I
(25), fig. 5
1886 Belemnites diceratianus Etallon – Loriol, pp. 37—38, pl. I,
figs. 1—4
1986a Hibolites semisulcatus (Münster) – Combémorel & Mariotti,
pp. 312—313, pl. 2, figs. 14—16 (cum syn.)
1986a Hibolites sp. – Combémorel & Mariotti, pp. 314, pl. 2,
figs. 17—20
pars 1990 Hibolithes semisulcatus (Münster) – Combémorel & Mariotti,
p. 213, pl. 2, figs. 5—8, 10
1995 Hibolithes semisulcatus (Münster) – Mariotti, p. 235,
pl. III, figs. 3—4
1999 Hibolithes semisulcatus (Münster) – Schweigert, pp. 3—4,
text-figs. 1, 3—4, pl. 1, figs. 1—4, pl. 2, figs. 1—6, pl. 3,
figs. 1—2, pl. 4, figs. 1—7, pl. 7, fig. 3 (cum syn.)
2002b Hibolithes semisulcatus (Münster) – Mariotti, pp. 218—219,
text-fig. 5.1—4
?2006 Hibolithes (Hemihibolites) ex gr. semisulcatus (Münster) –
Ippolitov, pp. 57, 59, 60, text-figs. 3a—g (morph A), 3d—z
(morph B)
2009 Hibolithes (gr.) semisulcatus (Münster) – Lukeneder, pl. 3, fig. G
M a t e r i a l : 31 near complete to fragmentary specimens
from Beds 77 to 25.
D e s c r i p t i o n : A slender rounded to dorso-ventrally
compressed hastate rostrum with pointed to obtuse centrally
placed apex. The alveolar area is characterized by a well-de-
veloped constriction towards the rostrum sollidum. Lateral
compression of the alveolar area is not always apparent, or
might be absent. The alveolus is shallow with a well devel-
oped fine but clear alveolar groove. The length of the groove
various, but might run well onto the rostrum sollidum. The
apical line is central.
R a n g e : Latest Oxfordian to middle Tithonian.
O c c u r r e n c e : Throughout the (sub-) Mediterranean
Tethys. Occasional occurrences outside this Province are not-
ed (Russian Platform; cf. Ippolitov 2006).
Genus: Subulibelus Riegraf, 1981
Subulibelus problematicus? Riegraf, 1981
1981 Subulibelus problematicus n. gen. et n. sp. Riegraf, pp. 99—100,
text-fig. 231, pl. 7, fig. 69
1981 Subulibelus cf. problematicus n. gen. et n. sp. – Riegraf,
pp. 101—102, text-fig. 232, pl. 7, fig. 70
M a t e r i a l : One near complete specimens from Bed 76
and three incomplete specimens from 75.
D e s c r i p t i o n : Small to medium sized very elongated,
very hastate specimens. In Bed 77 an incomplete specimen
shows a very shallow alveolus. No alveolar groove visible or
preserved.
R a n g e : Latest Oxfordian (?) to earliest Kimmeridgian.
O c c u r r e n c e : Latest Oxfordian (Ringsteadia pseudocor-
data Zone) and earliest Kimmeridgian (S. platynota Zone) of
southwest Germany (Riegraf 1981) and eventually Hungary.
429
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
Subulibelus? sp. or juvenile Hibolithes sp.
M a t e r i a l : Two specimens from Bed 75 and one from
Bed 76.
D e s c r i p t i o n : Very small specimens (< 15 mm) which
show a clear hastate sagital section. However, the potential
elongated proximal part of the rostrum is not preserved.
R a n g e : They occur in the ?Oxfordian-Kimmeridgian
boundary beds.
O c c u r r e n c e : Subulibelus is described from the latest
Oxfordian and the Oxfordian-Kimmeridgian boundary beds
of southwest Germany (Riegraf 1981).
Family: Duvaliidae Pavlow, 1914
Genus: Conobelus Stolley, 1919
Conobelus strangulatus (Oppel, 1865)
(Fig. 7.3,4)
1865 Belemnites strangulatus Oppel, p. 545
1868 Belemnites strangulatus Oppel – Zittel, pp. 35—36, pl. 1,
figs. 6—7
1890 Belemnites Orbignyi Duval-Jouve – Toucas, pp.587—588,
pl. XV, fig. 1
1890 Belemnites Orbignyi Duval-Jouve var. suborbignyi Toucas,
p. 588, pl. XV, fig. 2
?1922 Belemnites strangulatus Oppel – de Gregorio, p. 8, pl. I,
fig. 15
1942 Conobelus strangulatus Oppel – Mandev, pp. 51—52, pl. III,
figs. 3—4
(sic)1986a Rhopaloteuthis strangulatus (Oppel) – Combémorel &
Mariotti, pp. 307—308, pl. 1, figs. 12—15 (cum syn.)
(sic)1995 Rhopaloteuthis strangulatus (Oppel) – Mariotti, p. 233,
pl. II, fig. 11
(sic)1997 Rhopaloteuthis strangulatus (Oppel) – Combémorel, pl. 28,
fig. 13 [cast of HT]
non?1997 Rhopaloteuthis strangulata (Oppel) – Janssen, pp. 31—32, pl. 3,
figs. 3—4 [?non], nec figs. 5—6 [ = Conobelus incertus Weiss]
M a t e r i a l : Twelve complete to fragmentary specimens
from Beds 46 to 41.
D e s c r i p t i o n : Rounded to laterally flattened conobeloid
rostrum with well developed alveolar area. The dorsal and
ventral sides are near parallel, except for the apical part. The
alveolus is shallow and a faint but clear alveolar groove runs
well onto the rostrum sollidum. The apex is obtuse to mucro-
nate (in gerontic specimens) and shifted to the dorsal side;
this might not be clearly visible in (incomplete) immature to
juvenile specimens. The apical line is shifted to the ventral
side.
R a n g e : Middle Tithonian to (?) earliest Berriasian (top S.
semiforme—(?)early B. jacobi Zones).
O c c u r r e n c e : Mediterranean Tethys.
Conobelus cf. massimoi? (Mariotti, 2002)
?1873 Belemnites Beneckei nov. sp. Neumayr, p. 156 (16), pl. XXXI, fig. 1
M a t e r i a l : One incomplete, weathered specimen (Bed 70).
D e s c r i p t i o n : Stout, slightly hastate, medium to small
sized rostra with rounded cross-sections, a deep alveolar cav-
ity (almost halfway the rostrum). The alveolar groove could
not be seen. See Mariotti (2002b) for more specific details.
R e m a r k s : The specimen resembles the species de-
scribed by Mariotti (2002b). However, as the ontogeny is not
known the immature specimens from the same bed, and from
Bed 68, are gathered as Conobelus? sp.
R a n g e : Middle Late Kimmeridgian.
O c c u r r e n c e : Latest Early Kimmeridgian (Crusso-
liceras divisum Zone) of northern Italy, and possibly the up-
permost T. compsum to basal H. beckeri Zones of Hungary.
Genus: Duvalia Bayle, 1878
Duvalia cf. abeli (Vetters, 1905)
(Fig. 7.12,13)
?1894 Duvalia ensifer (Oppel) – Retowski, pp. 218—219, pl. XIV, fig. 1
1905 Belemnites Abeli Vetters, pp. 246—247, text-fig. 3
1986a Duvalia aesinensis nov. sp. – Combémorel & Mariotti,
pp. 304—305, pl. 1, fig. 4
1995 Duvalia aesinensis Combémorel & Mariotti – Mariotti,
pp. 230—231, pl. II, fig. 4
M a t e r i a l : One incomplete specimen from Bed 46.
D e s c r i p t i o n : The rostrum is characterized by an ex-
tremely laterally flattened rostrum sollidum with an angular
cross-section and with an expanded alveolar area (not pre-
served in our specimen). The alveolar opening is more or
less quadrangular, with a shallow alveolus and a short alveo-
lar groove (cf. Vetters 1905).
R a n g e : Middle Tithonian (top S. semiforme—(?)M. ponti
Zones).
O c c u r r e n c e : Crimea(?), Hungary, Italy and northern
Klippenbelt of Calcareous Alps.
Duvalia ensifer (Oppel, 1865)
(Fig. 7.14,15)
1865 Belemnites ensifer Oppel, p. 545
1868 Belemnites ensifer Oppel – Zittel, p. 36, pl. 1, figs. 9 [“Nor-
malform”], 10
?1868 Belemnites ensifer Oppel var. Zittel, pl. 1, fig. 11 [=?Duvalia esba]
?1875 Belemnites ensifer Oppel – Pillet & de Fromentel, pp. 14, 65,
pl. VIII, figs. 1—3 [ = ?Duvalia esba]
non1889 Belemnites ensifer Oppel – Sokolov, pp. 131—132, pl. II,
fig. 8a—c [ = Duvalia gr. lata]
non1894 Belemnites ensifer Oppel – Retowski, pp. 218—219, pl. XIV,
fig. 1[ = ?Duvalia abeli]
?1897 Belemnites ensifer Oppel – Roman, p. 279, pl. I, fig. 1
[ = ?Conobelus]
?1904 Belemnites ensifer Oppel – Schiller, p. 27 (133), text-fig. 3
[ = ?Conobelus]
non1922 Belemnites ensifer Oppel – de Gregorio, p. 8, pl. I, figs. 20
[ = ?Conobelus sp. indet.], 21 [ = ?“Pb.” gr. zeuschneri]
1935 Belemnites ensifer Oppel – Beregov, pp. 68 (19), 106 (56),
110 (60), pl. I, fig. 10
1986a Duvalia ensifer (Oppel) – Combémorel & Mariotti, pp. 303—
304, pl. 1, figs. 1—3
1990 Duvalia ensifer (Oppel) – Combémorel & Mariotti, pp. 210—
211, pl. 1, figs. 4—5
M a t e r i a l : Five incomplete specimens from Beds 46 to 44.
D e s c r i p t i o n : Typical duvaloid rostrum with (strong)
lateral compressed rostrum with elongated rounded cross-
430
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
sections and a dorsally orientated apex. The alveolus is mod-
erately deep and an alveolar groove is well developed, gen-
erally running well on to the rostrum sollidum. Lateral lines
(two parallel lines) may be visible on well-preserved speci-
mens. Juvenile to immature specimens do not show the same
outer-morphology as mature specimens.
R a n g e : Middle Tithonian (topmost S. semiforme—base S.
fallauxi Zones).
O c c u r r e n c e : Mediterranean Tethys.
Duvalia cf. esba (de Gregorio, 1885)
(Fig. 7.20,21)
?1868 Belemnites ensifer Oppel var. Zittel, pl. I(25), fig. 11
?1875 Belemnites ensifer Oppel – Pillet & de Fromentel, pp. 14, 65,
pl. VIII, figs. 1—3
1885 Belemnites esbus de Gregorio, p. 242
1886 Belemnites esbus de Gregorio – de Gregorio, p. 4, pl. 1,
figs. 12a—c
?1917 Belemnites (Duvalia) latus Blainville–Kilian & Révil, pl. XV, fig. 1
?1997 Duvalia sp. nov? spec. indet. Janssen, pp. 26—27, pl. 2, figs. 5—6
Material : One incomplete, juvenile specimen from Bed 32.
D e s c r i p t i o n : Typical duvaliid latatoid rostrum with a
laterally strongly compressed rostrum sollidum and a con-
stricted alveolar area. The alveolar groove is short and the al-
veolus is shallow. This species most probably belongs to the
group of belemnites around ensifer, but the overall shortage
of material does not permit more than morphologically com-
parison with the specimen depicted by de Gregorio. It ap-
pears to be much more laterally compressed as compared to
typical ensifer-species, with a more constricted alveolar area
and consequently a much shallower alveolus.
R a n g e : Late middle Tithonian—earliest Berriasian (top S.
fallauxi— (?) base B. jacobi Zones).
O c c u r r e n c e : Hungary, Italy, and possibly in Spain and
the Swiss Alps.
Duvalia tithonia (Oppel, 1865)
1865 Belemnites tithonius Oppel, p. 545
1868 Belemnites tithonius Oppel – Zittel, p. 37, pl. I, figs. 12—13
1889 Belemnites (Duvalia) Deeckei n. sp. Kilian, p.636, pl.
XXVI, fig. 5
1894 Belemnites tithonius Oppel – Retowski, pp. 221—222,
pl. XIV, figs. 3—4
non1922 Belemnites tithonius Oppel – de Gregorio, p. 8, pl. I, fig. 11
1983 Biplanidelus [sic!; recte Biplanibelus] tithonia (Zittel) –
Nerodenko, p. 42
(sic)1986b Duvalia tithonica (Oppel) emend. Zittel – Combémorel &
Mariotti, pp. 36—39, text-fig. 2 (cum syn.)
(sic)1990 Duvalia tithonica (Oppel) – Combémorel & Mariotti,
p. 211, pl. 1, fig. 6
(sic)1992 Pseudoduvalia tithonica (Oppel) – Barskov & Weiss, p. 74
(sic)1995 Duvalia tithonica (Oppel) em. Zittel – Mariotti, pp. 231—232,
pl. II, figs. 5—6, pl. III, fig. 10
(sic)1996 Duvalia tithonica (Oppel) – Eliáš et al., pp. 263, 267,
pl. IV, figs. 12—14
M a t e r i a l : One poorly preserved, incomplete juvenile to
immature specimen from Bed 18.
D e s c r i p t i o n: An atypical duvaliid rostrum characterized
by dorsal and ventral hollows. These are barely visible on
the dorsal side of this immature specimen. The rostrum
sollidum is laterally flattened, and the alveolar area is charac-
terized by a constriction. See Combémorel & Mariotti
(1986b) for more details.
R a n g e : Latest Tithonian—earliest Berriasian (Microcan-
thum—early Jacobi Zones).
O c c u r r e n c e : Mediterranean Tethys.
Genus: Produvalia Riegraf, 1981
Produvalia (?) sp. 1
M a t e r i a l : One incomplete specimen (Bed 68) which
shows the typical laterally flattened sub-rectangular duvaliid
cross-section. The section shows rounded lateral sides. The
ventral side is rounded to sub-angular while the dorsal out-
line is slightly flattened and a trace of a groove appears to be
visible. The alveolar cavity, just visible in the specimen, is
centrally placed.
R e m a r k s : The cross-section does remind of Produvalia,
more specific of the late Early to early Late Oxfordian
Produvalia neyrivensis (Favre) hence the generic attribution.
R a n g e : Late Kimmeridgian (earliest H. beckeri Zone).
Produvalia (?) aff. nicosiai? (Mariotti, 2002)
M a t e r i a l : Two incomplete specimens from the Beds 75
and 77 which show sub-rounded to sub-rectangular cross-sec-
tions. These sections are laterally compressed with near straight
lateral sides. Lateral incisions are faint but visible. Both the ven-
tral (slightly more inflated) as well as the dorsal side are round-
ed. The part from Bed 77 is best preserved (30 mm), and shows
an almost straight dorsal side and a slightly curved ventral side,
in lateral view. The ventral side narrows towards the alveolar
area. A dorsal groove is not visible and no alveolar part is pre-
served. The alveolar line is centrally placed. In this specimen
the cross-section near the alveolar region is visible. It shows a
rounded outline with weak dorso-lateral depressions. Overall,
the outline and lateral compression point to a duvaliid rostrum.
R e m a r k s : The general form is similar to Produvalia
nicosiai (Mariotti, 2002) but the anterior cross-section ap-
pears much more rounded and the dorsal side is less hastate.
R a n g e : Latest Oxfordian(?) to earliest Kimmeridgian (S.
platynota Zone).
Genus: Pseudobelus Auctorum non de Blainville, 1827
( = Pseudobelus s.l.; = ?genus novum)
“Pseudobelus” zeuschneri (Oppel, 1865)
(Fig. 7.8,9,16,17,24,25)
1865 Belemnites Zeuschneri Oppel, p. 545
1870 Belemnites Zeuschneri Oppel – Zittel, p. 28, pl. (I) 25, fig. 9
1889 Belemnites Zeuschneri Oppel – Sokolov, pp. 122—123,
pl. III, fig. 5
pars?1922 Belemnites Zeuschneri Oppel sp. aff. – de Gregorio, p. 8,
pl. I, figs. 14, 21
1986a Pseudobelus zeuschneri (Oppel) – Combémorel & Mariotti,
pp. 310—311, pl. 2, figs. 1—6 (cum syn.)
1990 Pseudobelus zeuschneri (Oppel) – Combémorel & Mariotti,
p. 212, pl. 1, figs. 11—14
431
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
1995 Pseudobelus zeuschneri (Oppel) – Mariotti, p. 234, pl. II,
fig. 10, pl. III, fig. 9 (cum syn.)
M a t e r i a l : Six incomplete to fragmentary specimens
from Beds 61 to 55.
D e s c r i p t i o n : Medium sized elongated duvaliid belem-
nites with vague lateral impressions, especially in the apical
area. The alveolus is relative deep and the alveolar groove
runs well on to the rostrum sollidum. The apex is shifted to
the dorsal side. Lateral sides are near parallel.
R e m a r k s : The lateral depressions which characterize
Pseudobelus s.s. appear to be different from the Tithonian—
earliest Berriasian Pseudobelus s.l. but are clearly compara-
ble to morphological features as can be observed on some
Produvalia and Duvalia. Combémorel & Mariotti (1986a:
p. 310) indicate that the original as depicted by Zittel (1870)
is not complete (anymore; lacking the alveolar part). Despite
that they do not mention it, in my opinion, the depth of the
lateral incisions is exaggerated.
R a n g e : Late Early Tithonian to earliest Late Tithonian
(probably Semiforme—Ponti Zones).
R a n g e : Early Tithonian (H. hybonotum—S. darwini Zones),
probably also in younger sediments.
O c c u r r e n c e : Mediterranean Tethys.
“Pseudobelus” ex gr. zeuschneri (Oppel, 1865)
(Fig. 7.5,6)
M a t e r i a l : Four incomplete to fragmentary specimens
from Beds 32 to 21.
D e s c r i p t i o n : Comparable to zeuschneri but with a well-
developed constriction between the alveolar area and the
rostrum sollidum. In the Hungarian material the alveolar ar-
eas are not preserved. Given the strong constriction, the
alveolus is most probably very shallow. Lateral lines are
very vague. The apex is pointed to mucronate, and shifted
towards the dorsal side. In general, these belemnites give the
impression of immature Duvalia.
R a n g e : Latest middle Tithonian—early Late Tithonian
(top S. fallauxi—base of M. microcanthum Zones).
O c c u r r e n c e : Hungary.
Acknowledgments: Prof. József Pálfy and PhD student
Márton Rabi (ELTE University, Budapest) are thanked for
their kind assistance in the field. Willem Bont (Naturalis,
Leiden) is thanked for some of the photographic work. This
work has been supported by the Hungarian Scientific
Research fund (OTKA) Grant: 68453 and received support
from the SYNTHESYS Project (http://www.synthesys.info/),
which is financed by European Community Research Infra-
structure Action under the FP6 Structuring the European
Research Area Program. The paper benefited from the com-
ments of reviewers, Drs Evgenij J. Baraboshkin (Moscow)
and Johann Schnyder (Paris).
References
Anderson T.F. & Arthur M.A. 1983: Stable isotopes of oxygen and
carbon and their application to sedimentologic and environ-
mental problems. In: Arthur M.A., Anderson T.F., Kaplan I.R.,
Veizer J. & Land L.S. (Eds.): Stable isotopes in sedimentary
geology. SEPM Short Course 10, 1—151.
Barskov I.S. & Weis A.F. 1992: On the ontogeny of some Early
Cretaceous belemnoids. Paleontological J. 26, 2, 70—85, 1 pl.
Bayle E. 1878: Fossiles principaux des terrains. Explication de la
carte géologique de France, série 4, 1 (Atlas), 176 pls.
Beregov R. 1935: Geology of the western part of Radomirsko. Rev.
Bulgar. Geol. Soc. 7, 2, 51(1)—113(63), 1 pl. (in Bulgarian).
Bettencourt V. & Guerra A. 1999: Carbon and oxygen isotope com-
position of the cuttlebone of Sepia officinalis: a tool for pre-
dicting ecological information? Mar. Biology 133, 651—657.
Blainville H.M. Ducratoy de 1827: Mémoire sur les bélemnites,
considérées zoologiquement et géologiquement. F.G. Levrault,
Paris, Strasbourg, 1—136, 5 pls.
Brand U. 1994: Morphochemical and replacement diagenesis of
biogenic carbonates. In: Wolf K.H. & Chilingarian G.V.
(Eds.): Diagenesis, IV. Elsevier, Amsterdam, 217—282.
Brand U. & Veizer J. 1980: Chemical diagenesis of a multicompo-
nent carbonate system: 1. Trace elements. J. Sed. Petrology 50,
1219—1286.
Cecca F. & Enay R. 1991: Les ammonites des zones
a
Semiforme et
a
Fallauxi du Tithonique de l’Ard
e
che (sud-est de la France):
stratigraphie, paléontologie, paleobiogeographie. Palaeonto-
graphica A219, 1—87, 10 pls.
Challinor A.B. & Hikuroa D.C.H. 2007: New Middle and Upper Ju-
rassic belemnite assemblages from West Antarctica (Latady
Group, Ellsworth Land): taxonomy and paleobiogeography.
Palaeontologia Electronica 10, 1, 6A, 1—29.
Combémorel R. 1997: Bélemnites. In: Cariou E. & Hantzperque P.
(Eds.): Biostratigraphie du Jurassique ouest-européen et médi-
terranéen: zonations parall
e
les et distribution des invert
e
brés et
microfossiles. Bull. Cent. Rech. Explor.-Production Elf Aquitaine,
Mém. 17, 157—162.
Combémorel R. & Mariotti M. 1986a: Les bélemnites de la carri
e
re
de Serra San Quirico (Province d’Ancona, Apennin central, Italie)
et la paléobiogéographie des bélemnites de la Téthys méditer-
ranéenne au Tithonique inférieur. Géobios 19, 3, 299—321.
Combémorel R. & Mariotti M. 1986b: First record of Duvalia ti-
thonica, a marker of Upper Tithonian, in Central Apennines.
Boll. Soc. Paleont. Ital. 25, 35—39.
Combémorel R. & Mariotti M. 1990: Taxonomic and biostrati-
graphic remarks on Tithonian belemnites from Sicily. Atti del
II Convegno Internazionale. Fossili, evoluzione, ambiente.
Pergola, 1987, 207—219.
Csontos L. & Vörös A. 2004: Mesozoic plate tectonic reconstruc-
tion of the Carpathian region. Palaeogeogr. Palaeoclimatol.
Palaeoecol. 210, 1—56.
Császár G. (Ed.) 1997: Basic lithostratigraphic units of Hungary.
Hung. Geol. Inst., Budapest, 1—114.
de Gregorio A. 1885: Tithonian fossils (Stramberger Schichten) of
the biancone of Rover
e
di Velo. Il Naturalista Siciliano 4, 1—6
(in Italian).
de Gregorio A. 1886: Fossiles tithoniques des Stramberg Schichten
du “Biancone”, “Rover
e
di Velo” des Alpes de Verone. Note
paléontologique. Ann. Géol. Paléont. 3, 1—8, 1 pl.
de Gregorio A. 1922: Monography on the Tithonian fossils of
“Casale di Sopra” (Busambra), preserved in my personal col-
lection (zone of Terebratula diphya e janitor). Ann. Géol.
Paléont. 36, 3—28, 12 pls. (in Italian).
Denys de Montfort P. 1808: Conchyliologie systématique, et classi-
fication méthodique des coquilles; offrant leurs figures, leur
arrangement générique, leurs descriptions caractéristiques,
leurs noms; ainsi que leur synomymie en plusieurs langues.
Tome 1. F. Schoell, Paris, 1—409, 100 pls.
Eliáš M., Martinec P., Reháková D. & Vašíček Z. 1996: Geology
è
è
è
è
è
è
à
à
432
FŐZY, JANSSEN and PRICE
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
and stratigraphy of the Kurovice Limestone and Tlumačov Marl
Formation at the Kurovice quarry (Upper Jurassic, Lower Creta-
ceous, Outer Western Carpathians, west Carpathians, Czech Re-
public). Věst. Čes. Geol. Úst. 71, 3, 259—276 (in Czech).
Enay R. & Geyssant J. 1975: Faunes Tithoniques des chaines bé-
tiques (Espagne méridionale). In: Colloque Limite Jurassique-
Crétacé, Lyon-Neuchatel, 1973. Mém. BRGM 86, 39—55.
Etallon A. 1862: Études paléontologiques sur la haut-Jura. Monog-
raphie du Corallien. Vertébrés, articulés, mollusques (Séances
des 8 janivier et 12 mars 1859). Mém. Soc. Émulation du Dé-
partement du Doubs, Ser. 3, 6 (for 1861), 53—227.
Favre E. 1877: La zone
a
Ammonites acanthicus dans les Alpes de la
Suisse et de la Savoie. Mém. Soc. Paléont. Suisse 4, 1—115, 9 pls.
Főzy I. & Meléndez G. 1997: Middle and Upper Oxfordian ammo-
nite succession from the Transdanubian Central Range and
from the Mecsek Mountains (Hungary): Biostratigraphy and
paleobiogeographic affinities. Comunicasiones IV Congreso de
Jurasico de Espana, Alcaniz 1997, 69—72.
Galácz A. 1975: Coupes bajociennes dans le Bakony Septentrional.
Földt. Közl. 105, 208—219 (in Hungarian).
Galácz A. & Vörös A. 1972: Jurassic history of the Bakony Moun-
tains and interpretation of the principal lithological phenomena.
Földt. Közl. 102, 2, 122—135 (in Hungarian).
Gemmellaro G.G. 1871: Paleontological studies on the “calcare a
Terebratula janitor” in the north of Sicily. Part 1. Lao, Palermo
1—56, 12 pls. (in Italian).
Géczy B. 1976: Les ammonites du Carixien de la Montagne du Ba-
kony. Akadémiai Kiadó, Budapest, 1—223.
Glover C.P. & Kidwell S.M. 1993: Influence of organic matrix on
the post-mortem destruction of molluscan shells. J. Geol. 101,
729—747.
Grabowski J., Haas J., Márton E. & Pszczółkowski A. 2010: Mag-
neto- and biostratigraphy of the Jurassic/Cretaceous boundary
in the Lókút section (Transdanubian range, Hungary). Stud.
Geophysica et Geodaetica 54, 1—26.
Haas J., Görög Á., Kovács S., Ozsvárt P., Matyók I. & Pelikán P.
2006: Displaced Jurassic foreslope and basin deposits of Di-
naridic origin in Northeast Hungary. Acta Geol. Hung. 49,
125—163.
Hikuroa D.C.H. 2004: The fauna and biostratigraphy of the Jurassic
Latady Formation, Antarctic Peninsula, Auckland. [Unpublished
Thesis University of Auckland], 1—399.
Ippolitov A.P. 2006: On the possible expression of sexual dimor-
phism in Hibolithes Montfort, 1808 from the Middle and Upper
Jurassic of European Russia. In: Barskov I.C. & Leonovoi T.B.
(Eds.): Contributions to current cephalopod research: morpholo-
gy, systematics, evolution, ecology and biostratigraphy. Mos-
cow, 8—10 November 2006. Dedicated to the 90
th
birthday of
the distinguished Russian researchers on fossil cephalopods
V.N. Shimansky and V.V. Drushchits. Russian Acad. Sci., Pa-
leont. Inst., Moscow, 57—60 (in Russian).
Janssen N.M.M. 1997: Mediterranean Neocomian belemnites. Part 1:
Río Argos sequence (province of Murcia, Spain): the Berria-
sian—Valanginian and the Hauterivian-Barremian boundaries.
Scripta Geol. 114, 1—55.
Kilian W. 1889: Études paléontologiques sur les terrains secondaires
et tertiaires de l’Andalousie. In: Bertrand M. & Kilian W.
(Eds.): Mission d’Andalousie. Mém. Acad. Sci. Inst. Nat. France
30, 2, 601—739, pls. 24—37.
Kilian W. & Révil J. 1917: Études géologiques dans les Alpes occi-
dentales. Contributions
a
la géologie des chaînes intérieures
des Alpes françaises. II. Description des terraines qui prennent
part
a
la constitution géologique des zones Intra-Alpines
françaises (Suite, syst
e
me jurassique). Mém. pour servir
a
l’Explication de la Carte Géologique détaillée de la France
1912, 1—273, pls. A—E, XII—XIX.
Loriol P. de 1886: Étude sur les mollusques des couches Coralli-
g
e
nes de Valfin (Jura), précedées d’une notice stratigraphique
par l’abbé E. Bourgeat. Partie I. Mém. Soc. Paléont. Suisse
XIII, 3, 1—120, pls. A—C, I—XI.
Lowenstam H.A. & Epstein S. 1954: Paleotemperatures of the post
Albian Cretaceous as determined by the oxygen isotope method.
J. Geol. 62, 207—248.
Lukeneder A. 2009: New biostratigraphic ammonite data from the Ju-
rassic/Cretaceous boundary at Nutzhof (Gresten Klippenbelt,
Lower Austria). Ann. Naturhist. Mus. Wien 110A, 313—329.
Mandev P. 1942: Géologie de la Zlatiška Planina et de ses avant-
monts dans le circuit du courant supérieur de la rivi
e
re Vit.
Rev. Bulgar. Geol. Soc. 13, 1 (for 1941), 1—71, pl. I—VI (in Bul-
garian with French abstract).
Mariotti N. 1995: Jurassic belemnites and aulacocerites from cen-
tral Italy. In: Farinacci A. (Ed.): Biostratigraphy of central Italy.
Stud. Geol. Camerti, Vol. Spec. 1994, parte A, 217—254 (in
Italian).
Mariotti N. 2002a: Upper Callovian—Middle Oxfordian belemnite
assemblages from the Monte Kumeta (Jurassic of western Sici-
ly, Italy). Boll. Soc. Paleont. Ital. 41, 13—35.
Mariotti N. 2002b: Systematics and taphonomy of an Early Kimmerid-
gian belemnite fauna from the Mediterranean Tethys (Monte
Nerone, Central Apennines, Italy). Geobios 35, 213—232.
Mariotti N. 2003: Systematics and taphonomy of an Early Kimmerid-
gian belemnite fauna from the Mediterranean Tethys (Monte
Nerone, Central Apennines, Italy). Géobios 36, 603—623 [cor-
rected reprint of 2002b].
Münster G. Graf zu 1830: Bemerkungen zum näheren Kenntniss der
Belemniten. F.C. Birner, Bayreuth, 1—18, 2 pls.
Nerodenko V.M. 1983: Early Cretaceous belemnites of the western
USSR. In: Starobogatov Ya.I. & Nesis K.N. (Eds.): Systematics
and evolution of cephalopod molluscs. Scientific Acad. USSR,
Zoological Inst., Leningrad, 42—43 (in Russian).
Neumayr M. 1873: Die Fauna der Schichten mit Aspidoceras
acanthicum. Abh. K.-Kön. Geol. Reichsanst. V, 141—257, pl.
31—43.
Nunn E.V., Price G.D., Hart M.B., Page K.N. & Leng M.J. 2009:
Terrestrial and marine carbon isotope signals from the Callov-
ian—Kimmeridgian (Late Jurassic) succession at Staffin Bay,
Isle of Skye, Scotland. J. Geol. Soc. London 166, 633—641.
Oloriz F. 1978: Kimmeridgian—lower Tithonian of the central part
of the Betic Cordilleras (Subbetic Zone) – Paleontology, bio-
stratigraphy. Univ. Granada, Thesis Doctoral 184, 1—758 (in
Spanish).
Oppel A. 1865: Die Tithonische Etage. Z. Deut. Geol. Gesell. 17,
535—558.
Pavia G. & Cresta S. (Eds.) 2002: Revision of Jurassic ammonites
of the Gemmellaro collections. Quad. Museo G.G. Gemmellaro,
Palermo 6, 1—408.
Pavlow A.P. 1914: Les cephalopodes du Jura et du Crétacé inférieur
de la Siberie septentrionale. Mém. Acad. Impéiale Sci. St.-
Pétersbourg, Ser. 8 – classe physico-mathématique, XXI, 4,
1—68, 18 pls. (in Russian).
Pillet L. & de Fromentel E. 1875: Description géologique et paléon-
tologique de la Colline de Lémenc sur Chambéry. Chatelain,
Chambéry, 1—135, 15 pls.
Price G.D. & Mutterlose J. 2004: Isotopic signals from late Juras-
sic—early Cretaceous (Volgian—Valanginian) sub-Arctic be-
lemnites, Yatria River, Western Siberia. J. Geol. Soc. London
161, 959—968.
Price G.D. & Rogov M.A. 2009: An isotopic appraisal of the Late
Jurassic greenhouse phase in the Russian Platform. Palaeo-
geogr. Palaeoclimatol. Palaeoecol. 273, 41—49.
Price G.D., Wilkinson D., Hart M.B., Page K.N. & Grimes S.T.
2009: Isotopic analysis of co-existing late Jurassic fish otoliths
à
è
à
à
è
à
è
433
AMMONITES, BELEMNITES AND STABLE ISOTOPES FROM UPPER JURASSIC BAKONY MTS (HUNGARY)
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2011, 62, 5, 413—433
and molluscs: implications for upper ocean water temperature
estimates. Geology 37, 215—218.
Retowski O. 1894: Die tithonischen Ablagerungen von Theodosia.
Ein Beitrag zur Paläontologie der Krim. Bull. Soc. Imperiale
Naturalist. Moscou 7, 206—301.
Riboulleau A., Baudin F., Daux V., Hantzpergue P., Renard M. &
Zakharov V. 1998: Évolution de la paléotempérature de eaux de
la plate-forme russe au cours du Jurassique supérieur. Comptes
Rendus de l’Académie des Sciences Série II, 326, 239—246.
Riegraf W. 1981: Revision der Belemniten des Schwäbischen Jura.
Teil 8. Palaeontographica (A) 173, 64—139, 5 pls.
Roman F. 1897: Recherches stratigraphiques et paléontologiques
dans le Bas-Languedoc (Th
e
se). Ann. Univ. Lyon 34, 1—345, 9
pls, 1 map.
Sarti C. 1984: Ammonite fauna and biostratigraphy of the central
Trentino region (Kimmeridgian-Tithonian), Northern Italy. Boll.
Soc. Paleont. Ital. 233, 473—514 (in Italian, English abstract).
Sarti C. 1993: Ammonite Fauna and biostratigraphy of the Venetian
Alp Kimmeridgian. Mem. Mus. Civi. Storia Naturale Verona,
1—145 (in Italian, English abstract).
Scherzinger A., Főzy I. & Parent H. 2010: Virgatosimoceras (Am-
monoidea, Simoceratidae) from the Lower Tithonian (Upper
Jurassic) – revision and value for correlation. Neu. Jb.
Paläeont., Abh. 256, 2, 195—212.
Schiller W. 1904: Geologische Untersuchungen im östlichen
Unterengadin. I. Lischannagruppe. Ber. Naturforsch. Gesell.
Freiberg im Breisgau 14, 107—180, 5 pls.
Schweigert G. 1999: Erhaltung und Einbettung von Belemniten im
Nusplinger Plattenkalk (Ober-Kimmeridgium, Beckeri-Zone,
Schwäbische Alb). Stuttgarter Beitr. Naturkunde (B; Geologie
und Paläontologie) 273, 1—35.
Shackleton N.J. & Kennett J.P. 1975: Paleotemperature history of
the Cenozoic and the initiation of Antarctic glaciation: Oxygen
and carbon isotope analyses in DSDP sites 277, 279 and 281.
In: Kennett J.P. & Houtz R.E. et al. (Eds.): Initial Reports of the
Deep Sea Drilling Project, 29. US Government Printing Office,
Washington, 743—756.
Sokolov V.D. 1889: Materials for the geology of the Crim. The Ti-
thonian of the Crim. Material for the Geology of Russia, XIII,
97—139, pl. II—IV (in Russian).
Stolley E. 1919: Die Systematiek der Belemnieten. Jahresbericht des
Niedersächsischen Geologischen Vereins 11 (for 1918), 1—59.
Tavera J.M. 1985: Upper Tithonian—Berriasian ammonites of the
Subbetic Zone (Betic Cordillera). Tesis Doctoral, Universidad
de Granada, 1—381 (in Spanish).
Toucas A. 1890: Étude de la faune des couches tithonique de
l’Ard
e
che. Bull. Soc. Géol. France, Sér. 3 18, 560—630, 6 pls.
Veizer J. 1983: Chemical diagenesis of carbonates: Theory and ap-
plication of trace element technique. In: Arthur M.A., Ander-
son T.F., Kaplan I.R., Veizer J. & Land L.S. (Eds.): Stable
isotopes in sedimentary geology. Soc. Econ. Paleont. Mineral-
ogist, Short Course Notes 10, 3.1—3.100.
Vetters H. 1905: Die Fauna der Juraklippen zwischen Donau
und Thaya. Beitr. Paläont. Österr.-Ungarns 17, 4 (for 1904),
223—259.
Vigh G. 1984: Die biostratigraphische Auswertung einiger Ammo-
niten-Faunen aus dem Tithon des Bakonygebirges sowie aus
dem Tithon-Berrias des Gerecsegebirges. Ann. Hung. Geol.
Inst. 67, 1—210.
Weissert H. & Channell J.E.T. 1989: Tethyan carbonate carbon iso-
tope stratigraphy across
the Jurassic—Cretaceous boundary:
an indicator of decelerated global carbon cycling? Paleoceano-
graphy 4, 483—494.
Weissert H. & Mohr H. 1996: Late Jurassic climate and its impact
on carbon cycling. Palaeogeogr. Palaeoclimatol. Palaeoecol.
122, 27—43.
Zittel K.A. von 1868: Die Cephalopoden der Stramberger Schichten.
Belemnites. In: Palaeontologische Studien über die Gren-
zschichten der Jura- und Kreide- Formation im Gebiete der
Karpathen, Alpen und Apenninen. Palaent. Mitt. Mus.
Koeniglichen Bayerischen Staates 2, 1, 33—38, 1 pl.
Zittel K.A. von 1870: Die Fauna der aeltern Cephalopoden fue-
hrenden Tithonbildungen. Palaeontographica, Suppl. 2, 2,
1—192, 15 pls.
è
è