GeolCarp_Vol62_No5_413

www.geologicacarpathica.sk

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

, NICO M.M. JANSSEN

and GREGORY D. PRICE

Department of Paleontology and Geology, Hungarian Natural History Museum, P.O. Box 137, H-1431 Budapest, Hungary;

fozy@nhmus.hu; semiformiceras@gmail.com

Geertekerkhof 14bis, 3511 XC Utrecht, The Netherlands; hibolithes@hotmail.com

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

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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.

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Table 3: Isotopic and elemental compositions of belemnite rostra from Lókút.

Belemnite species

Bed

number

(PDB)

(ppm)

Hibolithes cf. conradi*

Bed 16

0.9

1.4

371684

100

1890

240

Duvalia tithonia

Bed 18

–1.7

–1.5

355032

2026

1049

H. cf. conradi

Bed 18

–1.8

–0.9

359047

109

2317

972

Pseudobelus ex.gr. zeuschneri Bed 21

–1.0

–0.2

350100

1799

895

Belemnites sp.*

Bed 21

–0.3

–1.8

391288

1384

234

H. cf. conradi*

Bed 22

1.4

–0.4

356897

1615

270

Hibolithes sp.*

Bed 23

0.6

–1.2

385583

195

1963

179

208

H. cf. conradi

Bed 24

–1.6

–1.3

351484

2567

1055

Hibolithes sp. *

Bed 24

1.2

–2.6

361603

1993

140

525

Pseudobelus ex.gr. zeuschneri Bed 25

–2.3

–2.9

353975

2151

898

H. cf. semisulcatus*

Bed 25

–1.9

–2.2

349514

2231

104

794

H. cf. semisulcatus*

Bed 30

–0.8

0.4

359052

2758

125

1002

Pseudobelus ex.gr. zeuschneri Bed 32

–1.2

–0.9

351008

1860

955

Hibolithes sp.

Bed 32

–0.8

–1.1

362398

2431

995

H. cf. fellabrunnesis

Bed 32

–0.6

–1.3

362530

2503

864

Duvalia cf. ensifer*

Bed 32

0.4

388057

1939

591

470

H. cf. semisulcatus

Bed 35

–1.4

–1.6

361858

2525

1119

Hibolithes semisulcatus*

Bed 40

–1.2

–2.4

361213

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

1933

536

Hibolithes sp.

Bed 43

–1.8

–2.2

365793

1861

1020

Duvalia cf. ensifer

Bed 44

–0.2

–0.1

358767

2004

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

1365

115

334

Conobelus cf. strangulatus*

Bed 45

0.0

–1.0

358167

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

2253

939

Hibolithes semisulcatus*

Bed 45

1.3

–0.7

331695

152

2015

680

Hibolithes semisulcatus

Bed 46

–0.7

–1.1

339289

2286

771

Conobelus cf. strangulatus

Bed 46

–1.5

–1.2

351746

2166

1016

H. cf. semisulcatus

Bed 46

–0.9

–1.2

329436

2030

955

Conobelus cf. strangulatus*

Bed 46

–1.5

–0.1

343629

1792

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

1738

981

Duvalia cf. ensifer*

Bed 46

–0.5

0.3

361830

2171

1029

Hibolithes sp.*

Bed 47

–0.1

–1.3

356315

195

1834

869

458

Pseudobelus cf. zeuschneri

Bed 55

0.4

–1.2

356031

1827

989

H. cf. semisulcatus*

Bed 55

0.3

–1.6

363467

2851

1097

Hibolithes sp.

Bed 56

1.8

–1.4

381341

2627

932

Hibolithes sp.*

Bed 57

0.3

–0.5

346926

128

1927

684

713

Pseudobelus cf. zeuschneri

Bed 58

0.5

1.1

368994

1810

856

Hibolithes semisulcatus

Bed 58

1.4

–0.6

339196

1804

436

Hibolithes semisulcatus*

Bed 58

–0.4

–1.4

377196

2788

1131

Pseudobelus zeuschneri*

Bed 61

–0.7

–1.8

347528

1960

105

1067

Hibolithes sp. *

Bed 62

–0.1

–0.4

368284

2635

1077

Belemnites sp.

Bed 76

–0.3

0.5

365134

2328

855

Belemnites sp.

Bed 76

–0.6

0.2

369019

1626

918

Belemnites sp.

Bed 77

–0.5

0.3

370522

1500

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

and Mn

are more soluble under reducing conditions and thus available
for replacing Ca

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.

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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

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

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

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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

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

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

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

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-

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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.

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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-

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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

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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).

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