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Lower Toarcian Ammonitida fauna and

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 FRAGMENTA PALAEONTOLOGICA HUNGARICA 29, BUDAPEST, 2011

Lower Toarcian Ammonitida fauna and biostratigraphy of the Gerecse Mountains (Hungary)

by
Zoltn KOVCS

Abstract Stratigraphical and taxonomical problems of the Lower Toarcian (Jurassic) ammonite assemblages of the Gerecse Mts (NE
Transdanubian Range, Hungary) are discussed. The Submediterranean ammonite biostratigraphy with Dactylioceras tenuicostatum and
Harpoceras serpentinum Zones is established for the studied successions. A simplified stratigraphical scheme with two subzones (Harpoceras
serpentinum, Harpoceras falciferum) and three biohorizons (Hildaites striatus, Harpoceras pseudoserpentinum, Orthildaites douvillei) are recognized for the
Harpoceras serpentinum Zone. LowerMiddle Toarcian Ammonitida fauna is documented: species belonging to genera Phylloceras, Calliphylloceras,
Lytoceras, Dactylioceras, Mesodactylites, Nodicoeloceras, Catacoeloceras, Fontanelliceras, Polyplectus, Harpoceras, Ovaticeras, Hildaites, Orthildaites, Cingolites are
described and figured, a new species, Orthildaites becaudi n. sp. is introduced.

Key words Toarcian, Jurassic, Ammonitida, stratigraphy, Gerecse Mts, Hungary.

KOVCS, Z. (2011): Lower Toarcian Ammonitida fauna and biostratigraphy of the Gerecse Mountains (Hungary). Fragmenta Palaeontologica
Hungarica, 29: 148.

Introduction
This paper propounds a detailed taxonomical and strati- Toarcian Ammonitina fauna and biostratigraphy were
graphical revision of the Lower Toarcian ammonite assem- detailed by GCZY & SZENTE (2007); the taxonomy and
blages of the Gerecse Mts; as well as a comprehensive biostratigraphy of Upper ToarcianAalenian assemblages
ammonite biostratigraphy for the Toarcian Stage of the were dealt with by KOVCS & GCZY (2008), KOVCS
Gerecse successions is proposed. The ToarcianLower (2009, 2010), and KOVCS (in press); whereas the taxo-
Bajocian ammonite material comprises more than 15 000 nomical and paleobiogeographical problems of the lyto-
specimens, collected from six sections of the Gerecse Mts ceratid fauna was studied by KASSAI (2011). Besides the
(Nagy-Pisznice, Kis-Gerecse, Bnya-hegy AB, Tlgyht reinvestigation of the classic ammonite assemblages, two
AB; Figure 1) by the staff of the Geological Institute of other successions in the Gerecse Mts were recently treated.
Hungary between 1976 and 1982, and has been deposited in The mollusc fauna of the gastropod-rich ToarcianAalenian
the Natural History Museum of the Faculty of Science of beds of the Kis-Teke Hill Section was described by GALCZ
Etvs Lornd University, Budapest. & SZAB (2001) and GCZY et al. (2008). The newly exca-
vated Crocodile Section in the Nagy-Pisznice Hill yielded a
skeleton of a Steneosaurus specimen (SI et al. 2010). Its Toar-
cian biostratigraphy and the record of the ammonite fauna
were detailed by GALCZ et al. (in press). (Note: two sec-
tions were excavated on the Bnya-hegy: the MiddleUpper
Toarcian Ammonitina material of the A Section was detailed
in the mentioned papers, while the Lowerlower Middle
Toarcian fauna of the B Section is documented here.)
A comprehensive lithological, stratigraphical and paleo-
geographical examination of the Jurassic sediments of the
Gerecse Mts was accomplished by CSSZR et al. (1998);
and a geological review of the area was briefly offered by
GALCZ et al. (in press). Relatively complete Lower Jurassic
successions can be studied in the eastern part of the moun-
tains. The Upper Triassic Dachstein Limestone is overlain
by a sequence of deeper-water limestone that belongs to
Figure 1 Location of the ToarcianAalenian sections, the mainly pink or gray, well-bedded Pisznice Limestone of
examined in the Gerecse Mts. 35 m average thickness. It ranged to the Sinemurian and
the Lower Pliensbachian. It is overlain by the thinner (12
The first taxonomical, quantitative and paleobio- m) Trkbkk Limestone that represents the Upper Pliens-
geographical analyses of the Toarcian ammonites were bachian. The sedimentation was interrupted in the earliest
provided by GCZY (1984, 1985, 1990); the Middle Toarcian. The dark-grey or black clay (corresponding to the
Fragmenta Palaeontologica Hungarica 29, 2011
2 KOVCS, Z.
rkt Manganese Ore Formation of the Bakony Mts) that The carbonate-dominated succession is replaced by the
is typical of the Early Toarcian Oceanic Anoxic Event Ammonitico Rosso marl facies that developed in two sub-
cannot be traced in the studied sections except the Tlgyht facies in the Toarcian and Aalenian stages. The thin-bedded,
Quarry (SASVRI et al. 2009). (The thickness of Lower red nodular marl with variable carbonate and clay content
Toarcian strata of this section is appr. 165 cm. For some (Kisgerecse Marl) is typical of the Harpoceras serpenti-
insufficiency of the mentioned collection work, however, numGeczyceras speciosum Zones. It is followed by the
the presence of the Dactylioceras tenuicostatum Zone, and well-bedded, red, hard, nodular Tlgyht Limestone in the
the exact inner dimensions of the Harpoceras serpentinum Upper Toarcian. Both subfacies are rich in ammonites;
Zone are uncertain at this section. Its ammonite assemblage however, the fauna mainly consists of poorly to moderately
and stratigraphy need reinvestigation in another paper.) preserved internal moulds only.
Lower Toarcian ammonite biostratigraphy of the Gerecse Mountains
The stratigraphical analysis of this paper is based on studies relating the SubmediterraneanMediterranean areas
the standard ammonite zonation of the Toarcian Stage, summed up by VENTURI & BILOTTA (2008), METODIEV
which was put forward by ELMI et al. (1997), and reinter- (2008), BILOTTA et al. (2010), and COMASRENGIFO et
preted by PAGE (2003), and on the achievements of the al. (2010).
Dactylioceras tenuicostatum Zone
Index: Dactylioceras (Orthodactylites) tenuicostatum (YOUNG & Mts. Bed 113b, with first occurrence (FO) of Hildaites murleyi
BIRD, 1822). in the characteristic red Kisgerecse Marl facies, is considered
Author: BUCKMAN (1910). as belonging to the Harpoceras serpentinum Zone. The
In the Gerecse Mts, the zone was first identified by dactylioceratids of this bed (Orthodactylites aff. directum,
GCZY (1984, 1985), based on the presence of a Fontanelli- Mesodactylites aff. annulatiformis) also indicate the latter zone
ceras cf. fontanellense specimen in the lowermost bed of the (Table 1).
Kis-Gerecse Section. This locality was designated by KONDA In the Nagy-Pisznice Section, Harpoceras serpentinum
(1986) as the type section of the Kisgerecse Marl Formation, Zone strata (Beds 145133) are deposited with hiatus on
as well as briefly discussed by JENKYNS et al. (1991) and massive limestone beds belonging to the Pliensbachian
CSSZR et al. (1998). According to the authors, the Dacty- Trkbkk Limestone (the columnar section was figured by
lioceras tenuicostatum Zone is restricted to the Kis-Gerecse CSSZR et al. 1998, fig. 8). The Kisgerecse Marl appears in
Section with thin beds of 15 cm. This result is to be com- Bed 145 without yielding any Ammonitina specimens. In the
pleted, due to the revision of the ammonite assemblage. Bnya-hegy B Section the Dactylioceras tenuicostatum Zone
Two beds were recognized by GCZY (l.c.) as being part of is also missing. The top of the hardground of Trkbkk
the zone (Beds 113112), however, sedimentological diffi- Limestone beds are overlaid by the Kisgerecse Marl of the
culties occur. According to KONDA, the hard, nodular lime- Harpoceras serpentinum Zone with hiatus. Some poorly
stone layers of the Kisgerecse Marl are characterised by con- preserved ammonites (Canavaria sp. indet. [Plate V: 6],
densation, so the lithological boundary coincides with zonal Protogrammoceras ? sp.) came from Beds 1615; they are
boundary (1986). In contrast with this opinion, the litholo- thought to represent the Emaciaticeras emaciatum Zone
gical analysis shows Bed 113 being mixed, so it needs to be (Table 2). The chronostratigraphic boundary of this sequence
divided. Consequently, only bed 113a is considered here as resembles those described by GALCZ et al. (2008) from
representing the Dactylioceras tenuicostatum Zone. This Bakonycsernye. As the Bnya-hegy B Section was destroyed
bed yielded Fontanelliceras in a hard, brownish limestone by the quarrying, more detailed comparison cannot be
matrix, which is unknown in other localities of the Gerecse achieved.

Harpoceras serpentinum Zone

Index: Harpoceras (Harpoceras) serpentinum (SCHLOTHEIM, nean Province (e.g. Italy, Greece, Austria, Hungary, southern
1813) Spain, North Africa) was dealt with by GUEX (1973), ELMI
Author: OPPEL (1856) et al. (1989), and JIMNEZ & RIVAS (1991, 1992) (Figure 2).
Two regions were recognized in Europe by ELMI et al. It shows a less certain zonation with the Hildaites levisoni
(1997), and PAGE (2003) in the second Toarcian zone. The Zone including Hildaites levisoni and Harpoceras falciferum
standard NW European Province (e.g. southern England, Subzones. Without general agreement on the index fossils
France, Germany, the Iberian Range of Spain, Portugal) is the horizon-level subdivision and the lower boundary of the
characterized by the Harpoceras serpentinum Zone includ- zone remained debated. As genera Eleganticeras and Harpo-
ing two well-defined subzones with four horizons. This ceras are rare in Tethyan areas, the base of the zone is thought
stratigraphic pattern met with a wide acceptance in the litera- to coincide with the FO of genus Hildaites; while the upper
ture, while the region was reinterpreted as Submediterranean boundary is generally defined by the FO of Hildoceras sublevi-
Province by PAGE (l.c.) distinguishing the Subboreal Province soni. This scheme is accepted in the literature (ELMI 1986,
(northern Britain). The zonal subdivision of the Mediterra- BENSHILI 1989, FARAONI et al. 1994, PARISI et al. 1998,
Fragmenta Palaeontologica Hungarica 29, 2011
 Lower Toarcian Ammonitida 3
Table 1 Ammonitida distributions of Lower Toarcian of the Kis-Gerecse Section. D = Domerian, E = Emaciaticeras
emaciatum Z., T = Dactylioceras tenuicostatum Z., Serp. = Harpoceras serpentinum Subz.

Table 2 Ammonitida distributions of Harpoceras serpentinum Hildoceras bifrons Zones of the Bnya-hegy B
Section. D = Domerian, E = Emaciaticeras emaciatum Z., Douv. = Orthildaites douvillei H.

Fragmenta Palaeontologica Hungarica 29, 2011

4 KOVCS, Z.
Table 3 Ammonitida distributions of Harpoceras serpentinum Zone of the Crocodile Section. D = Domerian.

Table 4 Ammonitida distributions of Harpoceras serpentinum Zone of the Nagy-Pisznice Section. D = Domerian, E
= Emaciaticeras emaciatum Z.

EL HAMMICHI et al. 2009), however, the definition of the aside Harpoceratinae species as zonal indices raises problems
lower boundary was questioned by MACCHIONI (2002b) with respect to the correlatability between the NW
and MACCHIONI & CECCA (2002). Accordingly, putting European and Mediterranean ammonite biostratigraphy,

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 5
because the FO of genus Hildaites seems to precede the FO the dominance of the former two genera in NW European
of genus Harpoceras in Mediterranean successions (see GOY areas. Consequently, the equivalence of the Harpoceras
et al. 1988; ELMI et al. 1989; JIMNEZ et al. 1996). On the serpentinum and Hildaites levisoni Zones seems to be
other hand, apart from both Hildaites striatus and H. wrighti, acceptable in the literature (see PAGE 2003, 2004; OGG
which are thought to appear in the uppermost Dactylioceras 2004, BCAUD 2006, VENTURI & BILOTTA 2008, METODIEV
tenuicostatum Zone of British, Spanish, Italian and North 2008, BILOTTA et al. 2010). Nevertheless, the subdivisions of
African localities, the coeval FO of Harpoceras (or genus Ele- the zone are controversial in the papers that concern with
ganticeras) and other species of genus Hildaites seems to be the Tethyan area: various subzones and horizons were
evidenced in all three provinces in question, effectively with employed in different articles (Figure 2).

Figure 2 Correlation chart of the ammonite stratigraphy of the second Toarcian zone in the Submediterranean and
the Mediterranean Provinces.

The second Toarcian zone in the Gerecse Mts was first 14), and the internal moulds of the ammonites with cover
treated by GCZY (1984; 1985) as Hildaites serpentinus of black manganese crusts are common in three sections
Zone. In this paper, the formerly collected ammonite faunas (Kis-Gerecse: Beds 113b112, Bnya-hegy B: Beds 1410,
of the Kis-Gerecse, Nagy-Pisznice and Bnya-hegy AB Nagy-Pisznice: Beds 144141, 134133.)
Sections are reinvestigated, and completed with the records The Gerecse ammonite assemblages represent a charac-
of the Crocodile Section. (Only two beds are mentioned teristic Mediterranean faunal composition in the Middle
here from the the Bnya-hegy A Section: Bed 47 yielded no Upper Toarcian, with the dominance of suborders Phyllo-
Ammonitina, while Bed 46 belongs to the Orthildaites douvillei ceratina and Lytoceratina. The fauna is closely allied to those
horizon of the Harpoceras serpentinum Zone.) The average of Italy and Greece; however, typical NW European taxa of
thickness of the latter zone is approximately 120 cm in the Ammonitina were also recognized (GCZY 1990; KOVCS
studied sections. The sequences suggest continuity: the litho- 2009). The twofold affinity emerges in the Lower Toarcian
facies (Kisgerecse Marl Formation) is uniform; the thick- material as well. The low Ammonitina ratio is a clear
nesses of the zone are closely related in three well-exposed Mediterranean feature while occurrences of some Harpoceras
sections (Figure 3); the faunas are markedly similar (Tables species, e.g. H. strangewaysi, H. serpentinum, and H. pseudoserpen-
Fragmenta Palaeontologica Hungarica 29, 2011
6 KOVCS, Z.
tinum, which are typical of the SubborealSubmediterranean literature. Provisional stratigraphic approaches were recently
regions, are noteworthy. Similarly, genus Hildaites is also offered by BILOTTA et al. (2010), and SASSAROLI & VEN-
characterised by both SubborealSubmediterranean species TURI (2010), based on immensely diversified faunas of new
(Hildaites wrighti, H. murleyi, H. levisoni, H. forte, H. serpentini- Italian exposures (Figure 2). The lower boundary of the
formis), and species, which are typical of the Mediterranean Levisoni/Striatus Zone of Marconessa Section 2 was defined
Province (H. striatus, H. crassus, H. cf. undicosta, H. aff. by the first occurrence of Harpoceras gr. serpentinum and H. gr.
pseudolevisoni). This general feature of the Gerecse material maghrebensis GUEX, but the stratigraphical role of subfamily
confirms the new paleobiogeographical results submitted by Harpoceratinae in the further subdivison was replaced by
DERA et al. (2011). Accordingly, apart from the areas of SW frequent Hildaites species. Taking into account the altering
Tethys and the ApenninesIonian islands, remarkable inter- dominance of Hildaites striatus, H. undicosta and Orthildaites
provincial faunal mixing was characteristic for the Tethyan douvillei, three main bioevents were recognized in the second
Realm between the Dactylioceras tenuicostatum and the Toarcian zone (BILOTTA et al. 2010, fig. 3; SASSAROLI &
Hildoceras bifrons Zones. VENTURI 2010, fig. 2, tab. 1). Though considerable differ-
ences might be demonstrated between the Italian and Hun-
garian successions compared here (degree of condensation,
diversity, etc.), the Gerecse ammonite material seems to be
closely related to the Italian one, therefore the new biostrati-
graphical pattern appears to be applicable for the highly
condensed Gerecse sequences.
The local reconciliation of Mediterranean and standard
subdivisions proposed here is based on the conceptions of
METODIEV (2006, 2008) and BILOTTA et al. (2010); and
resulted in a slightly simplified biostratigraphic scheme with
two subzones and three biohorizons (Figure 4). Harpoceras
index species made possible to establish the Submediterranean
subzonation, which provides correlatibility with other regions.
The reason of use of Harpoceras serpentinum Subzone pro-
pounded by METODIEV (2006) is the lack of two standard
subzonal indices: Eleganticeras elegantulum (YOUNG & BIRD)
and E. exaratum (YOUNG & BIRD) in the fauna. (The latter
species is known from the Mecsek Mts in Hungary, but this
area belongs to the NW European Province). The Harpo-
ceras serpentinum Subzone is considered here as corres-
ponding to the standard Eleganticeras elegantulum Subzone.
On the other hand, the ammonites known from beds of the
Harpoceras serpentinum Subzone did not allow the Sub-
mediterranean horizon-level subdivision (Eleganticeras elegantu-
lum, Harpoceras strangewaysi), but demonstrated the relative
abundance of the Mediterranean horizon marker species
Hildaites striatus (see JIMNEZ 1986). In NW European
Figure 3 Zonal and horizon subdivision of the Lower localities Hildaites wrighti is typical of the lower, and H.
Toarcian of the studied sections. (The inner dimension
of the Kis-Gerecse Section diagram is estimated, as the thicknesses
subserpentinus is typical of the upper part of the zone
of beds were not recorded.) (BCAUD 2006, GOY et al. 2010), while in the Gerecse
sections these species co-appear in the condensed strata of
Based on Harpoceras index species, the Gerecse sequences the Hildaites striatus biohorizon. Due to the co-occurrence of
can be correlated with the Submediterranean zonation, and Harpoceras strangewaysi, H. serpentinum, Hildaites striatus, H.
the Harpoceras serpentinum Zone is to be introduced, murleyi and H. subserpentinus, the Hildaites striatus biohorizon is
instead of the Hildaites levisoni Zone which was proposed correlatable with the upper part of the standard Eleganticeras
for Mediterranean areas. The base of the zone in these suc- elegantulum Subzone without any more precise zonation. The
cessions shows close affinities to those of Tethyan localities: lower boundary of the Harpoceras serpentinum Subzone is
Hildaites appears before Harpoceras in three sections (Kis- defined by either the co-appearance of genera Harpoceras and
Gerecse, Bnya-hegy B, Crocodile). It is only the Nagy-Pisz- Hildaites (Nagy-Pisznice), or the FO of Hildaites murleyi (Kis-
nice Section where the FO of both genera is known from Gerecse and Bnya-hegy B) or H. subserpentinus (Crocodile
the same bed (Table 4). The examination made it obvious Section). The Harpoceras falciferum Subzone (HAUG 1885)
that the standard Submediterranean biostratigraphy can be also yielded a mixed MediterraneanSubmediterranean
adapted, however, with minor modification. Some sort of material with some excellent index taxa, so the introduction
integration of the SubmediterraneanMediterranean subdi- of Submediterranean horizons seems well-established. The
visions seems to be necessary by adopting new results of the lower boundary of the Harpoceras pseudoserpentinum biohorizon

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 7
(GABILLY 1976) coincides with the FO of Harpoceras falciferum, from the fauna, while genus Harpoceras shows an extended
H. pseudoserpentinum or H. mediterraneum. This horizon is well- distribution with two frequent taxa (H. mediterraneum and H.
developed in four sections (Figure 3), and characterised by the subexaratum) in the Hildoceras bifrons Zone as well.
dominance of Harpoceras and Hildaites; however, the boundary On the basis of the preceding comprehensive treatments
between the two subzones is uncertain in the Crocodile of the Gerecse ammonite material (see Introduction), and
Section (Table 3). The Orthildaites douvillei biohorizon (GOY & the stratigraphical achievement of this paper, the following
MARTNEZ 1990) is defined by the FO of the nominal index summary of ammonite biostratigraphy is proposed here for
species. At the top of this horizon genus Hildaites disappears the Toarcian Stage of the Gerecse successions (Figure 4).

Figure 4 Summarized ammonite biostratigraphy of the Toarcian Stage of the Gerecse Mts.

Systematic paleontology
The ammonite material shows a clear Tethyan character
with the dominance of suborders Phylloceratina and Lyto-
ceratina. Strata of the Harpoceras serpentinum Zone yielded
more than 1100 Ammonitida specimens of which 56%
represent Phylloceratina, 13% Lytoceratina, and 31% sub-
order Ammonitina (the statistical summary is based on
both the unpublished preliminary reports of the collection
work in the Kis-Gerecse, Nagy-Pisznice and Bnya-hegy
Sections (GCZY 1977, 1978, 1981), and the records of the
Crocodile Section). 240 Ammonitina specimens have been
identified at generic level. They represent two superfamilies
(Eoderoceratoidea, Hildoceratoidea) with two families and Figure 5 Familysubfamily level proportions of the
five subfamilies (Dactylioceratidae: Dactylioceratinae, Nodi- Gerecse Ammonitina fauna.
coeloceratinae; Hildoceratidae: Hildoceratinae, Polyplect-
inae, Harpoceratinae). Only the Lower Toarcian ammonite material of the
The superfamily Hammatoceratoidea, which appears in Gerecse Mts is analysed here; for Middle to Upper Toarcian
Lower Toarcian strata of Apenninic localities, is missing range of Ammonitina, see the above mentioned papers. As
from the material; it can be recorded only from the Hildo- some species has been discussed in detail in the literature, it
ceras bifrons Zone of the Gerecse assemblages (KOVCS seems sufficient to submit only selected synonymy. The
2009). The proportion of the Ammonitina faunal com- well-documented presence of species in both Submedi-
position is shown on Figure 5, and the stratigraphic terranean and Mediterranean European localities is indicated
distributions of Ammonitida are shown on Tables 14. by Europe in the paragraph Distribution.

Fragmenta Palaeontologica Hungarica 29, 2011

8 KOVCS, Z.
Abbreviations in the text E = external lobe, L = lateral lobe, ES = external saddle, LS = lateral saddle, D = diameter, H = whorl-height,
W = whorl-width, U = umbilical-width. Abbreviations in the Plates The figured specimens are coated with ammonium chloride, and
shown in natural size. The last septal suture is marked by an arrow. Abbreviations of measurements: D diameter, H whorl-height (H/D%), W
whorl-width (W/H%), U umbilical-width (U/D%), L length of fragment.

Suborder Phylloceratina ARKELL, 1950

Superfamily Phylloceratoidea ZITTEL, 1884
Family Phylloceratidae ZITTEL, 1884
Subfamily Phylloceratinae ZITTEL, 1884
Genus Phylloceras SUESS, 1865
Phylloceras heterophyllum (SOWERBY, 1820)
(Plate I: 14)
1820: Ammonites Heterophyllus SOWERBY, p. 119, pl. 266.
1845: Ammonites heterophyllus SOWERBY DORBIGNY, p. 339, pl. 109, figs 24.
1934: Phylloceras heterophyllum (SOWERBY) NEGRI, p. 113, text-fig. 5, pl. 10, fig. 1.
1961: Phylloceras heterophyllum (SOWERBY) KRYMHOLTS, p. 19, pl. 1, figs 12.
1963: Phylloceras heterophyllum (SOWERBY) ZANZUCCHI, p. 107, pl. 13, fig. 2.
1975: Phylloceras heterophyllum (SOWERBY) DEZI & RIDOLFI, p. 14, figs 1921.
1994: Phylloceras heterophyllum (SOWERBY) JOLY, p. 97, text-fig. 29, pl. 32: 46.
2000: Phylloceras heterophyllum (SOWERBY) JOLY, p. 43, text-figs 7278, pl. 5, figs 12, pl. 6, figs 13 (cum syn.).
2007: Phylloceras heterophyllum (SOWERBY) RULLEAU, p. 45, fig. 10/1, pl. 1, fig. 1.

Material from the H. serpentinum Zone 23 internal as typical of the Lower Toarcian Gerecse assemblages: Ph.
moulds of different state of preservation. heterophyllum (SOWERBY) and Ph. doderleinianum (CATULLO).
Description Large, involute, moderately compressed The latter differs by markedly wider whorl-section and
form with elliptical whorl-section and very narrow umbili- asymmetrical L lobe. Our Ph. heterophyllum specimens agree
cus. The flanks are somewhat convex, the venter is narrow, well with the type (SOWERBY 1820, pl. 266, refigured by
moderately low and rounded. No whole body chamber is ARKELL 1957, fig. 218.5), and the specimens figured in the
preserved. The ornamentation of well-preserved specimens literature in style of coiling, ornamentation and suture
consists of fine, dense, straigth and rectiradiate riblets construction. The species shows slight variability in width
passing over the venter. The suture-line consists of short E, (W/H: 5058%), but is a characteristic compressed form of
relatively wide L, and four auxiliary lobes. The ES is genus Phylloceras in the H. serpentinum H. bifrons Zones.
diphyllic, the lateral saddles are triphyllic, and the LS1 is Distribution Europe, North Africa, Caucasus,
higher than the ES. northern Russia. Hungary: Bakony Mts; Gerecse Mts: H.
Remarks Two species are recognized in this paper serpentinum H. bifrons Zones: all studied Gerecse sections.

Phylloceras doderleinianum (CATULLO, 1853)

(Plate I: 56, Plate II: 56)
1853: Ammonites Doderleinianus CATULLO, p. 19, pl. 1, fig. 3.
1934: Phylloceras Doederleinianum (CATULLO) NEGRI, p. 127, text-fig. 10, pl. 11, figs 25.
1963: Phylloceras doderleinianum (CATULLO) ZANZUCCHI, p. 108, pl. 13, figs 1, 4 (cum syn.).
v 1967a: Phylloceras doderleinianum (CATULLO) GCZY, p. 16, text-fig. 10, pl. 1, fig. 4, pl. 63, fig. 8.
1970: Phylloceras doderleinianum (CATULLO) PANNUZI, p. 60, pl. 2. figs 17.
1975: Phylloceras doderleinianum (CATULLO) DEZI & RIDOLFI, p. 14, figs 1618.
2001: Phylloceras doderleinianum (CATULLO) VENTURI & FERRI, p. 59.
2010: Phylloceras doderleinianum (CATULLO) VENTURI et al., p. 126.

Material from the H. serpentinum Zone 18 internal Ornamentation is not characteristic, however, narrow and
moulds of different state of preservation. shallow dorsolateral constrictions may appear. The suture-
Description Medium-size, involute form with wide- line consists of short E, relatively wide, asymmetrical L,
oval to rounded subrectangular whorl-section. The umbili- and four auxiliary lobes. The ES is diphyllic, the lateral
cus is narrow, the flanks are convex, the venter is low, saddles are triphyllic.
broad and rounded. No whole body chamber is preserved. Remarks The revision of the type was accomplished

Explanation to Plate I
12 Phylloceras heterophyllum (SOWERBY) (204.2011), Nagy-Pisznice, Bed 134, O. douvillei horizon, D: 96, H: 56 (58.3%), W: 30
(53.5%), U: 5 (5.2%).
34 Phylloceras heterophyllum (SOWERBY) (205.2011), Kis-Gerecse, Bed 103, H. pseudoserpentinum horizon, D: appr. 86, H: 50
(58%), W: 22 (44%), U: ?.
56 Phylloceras doderleinianum (CATULLO) (206.2011), Nagy-Pisznice, Bed 136, H. pseudoserpentinum horizon, D: 132, H: 76
(57.5%), W: 40 (52.6%), U: 7 (5.3%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 9
Plate I

Fragmenta Palaeontologica Hungarica 29, 2011

10 KOVCS, Z.
by MENEGHINI (1875, 18671881); the species differs from which were regarded as specific feature of Ph. loczyi (GCZY
Ph. heterophyllum in lower venter and wider whorls (W/H: 1967a: 18), appear on Ph. doderleinianum specimens figured
5966%). As it shows moderate variability in the width of from Italy (MAVIGLIA 1940, p. 14, fig. 9; ZANZUCCHI
the whorl-section and in the ornamentation, similar wide 1963, pl. 13, fig. 4; VENTURI 1982, p. 35, fig. 25; VENTURI
suboval forms with identical stratigraphic range and similar & FERRI 2001, p. 59).
suture-line (asymmetrical L lobe) are considered here as Distribution Italy, Greece, Albania, Montenegro,
synonyms of Ph. doderleinianum: Ph. gajarii PRINZ, Ph. borni Austria. Hungary: Bakony Mts; Gerecse Mts: H.
PRINZ, Ph. bckhi PRINZ, Ph. loczyi PRINZ, Ph. mitzopoulosi serpentinum H. bifrons Zones: all studied Gerecse
NEGRI. The weakly developed dorsolateral constrictions sections.

Subfamily Calliphylloceratinae SPATH, 1927

Genus Calliphylloceras SPATH, 1927
Calliphylloceras capitanii (CATULLO, 1847)
(Plate II: 12, Plate III: 56)
1847: Ammonites Capitanii, nob. CATULLO, p. 5, pl. 12, fig. 4ac.
1853: Ammonites Capitanei CAT. CATULLO, p. 38, pl. 4, fig. 4ac.
18671881: Phylloceras Capitanei CAT. MENEGHINI, p. 94, pl. 18, figs 46.
non 1911: Phylloceras Capitanei CAT. VADSZ, p. 64, text-fig. 17.
1936: Phylloceras Capitanioi (CATULLO) NEGRI, p. 53, pl. 12, figs 4, 5, 6; 1936, pl. 1, figs 1, 2.
1970: Calliphylloceras capitanioi (CATULLO) PANNUZI, p. 64, pl. 3, figs 16 (cum syn.).
2001: Calliphylloceras capitanioi (CATULLO) VENTURI & FERRI, p. 65.
2002a: Calliphylloceras capitanei (CATULLO) MACCHIONI, p. 46, fig. 15.

Material from the H. serpentinum Zone 13 internal specific features, however, without any consensus. Moreover,
moulds of different state of preservation. possibility of identity of the two taxa was raised by
Description Medium-size, involute, compressed MACCHIONI (2002a), with priority of CATULLOs species.
form with narrow oval whorl-section. The umbilicus is Comparing the two types (CATULLO 1847, l.c., and Ammon-
narrow, the flanks are slightly convex, the venter is rounded. ites calypso, DORBIGNY 1845, pl. 110, figs 13), and the speci-
No whole body chamber is preserved. The ornamentation mens figured by MITZOPOULOS (1930, pl. 2, figs 1, 2, 4),
is characterized by prorsiradiate, narrow, shallow, concave RAMACCIONI (1939, pl. 10, fig. 21, pl. 11, fig. 3), MAXIA
and curved constrictions, which become shallower on the (1943, pl. 1, figs 2, 3, 4), ZANZUCCHI (1963, pl. 13, figs 6, 7,
ventrolateral part, and pass over the venter. The specimens 12, 13) and VENTURI et al. (2010: 123, 132); the shape and the
bear 67 constrictions at diameter 60. The suture-line consists number of the constrictions are recognized as specific
of narrow E with half-length of the broader, long, symmet- morphological features in this paper. Consequently, C. nilssoni
rical L lobes, diphyllic ES, triphyllic LS1, and diphyllic is characterized by more pronounced, slightly more distant
LS2LS7. and less curved constrictions. The Gerecse specimens well
Remarks As the morphology, ornamentation and agree with the type (CATULLO 1847, pl. 12, fig. 4ac). C.
suture construction of the species are markedly similar to bicicolae (MENEGHINI) and C. beatricis (BONARELLI) are also
those of C. nilssoni (HBERT), many authors detailed the similar forms; however, the former possesses wider whorls,
differences between the two forms (e.g. PRINZ 1904; NEGRI and is typical of the Late Pliensbachian, while the latter is a
1936; ZANZUCCHI 1963; KOLLROVANDRUSOV 1966; smaller form with somewhat wider and straight constrictions.
JOLY 1994; MACCHIONI 2002a). Shape of the whorl-section, Distribution Italy, Greece, Austria, Albania, Algeria,
width of the umbilicus, number of the constrictions, number (?)Turkey. Gerecse Mts: H. serpentinum H. bifrons Zones:
of the lateral lobes, or stratigraphic range were mentioned as all studied Gerecse sections.

Calliphylloceras nilssoni (HBERT, 1866)

(Plate II: 34)
1866: Ammonites nilssoni HBERT, p. 527, fig. 3.
1927: Phylloceras Nilssoni (HBERT) SCHRDER, p. 127, pl. 7, figs 7ac, 8.

Explanation to Plate II
12 Calliphylloceras capitanii (CATULLO) (207.2011), Nagy-Pisznice, Bed 133, O. douvillei horizon, D: 100, H: 53 (53%), W: 24
(45.2%), U: 7 (7%).
34 Calliphylloceras nilssoni (HBERT) (210.2011,) Kis-Gerecse, Bed 100, H. sublevisoni horizon, D: 53, H: 28 (52.8%), W: 18
(64.2%), U: 4 (7.5%).
56 Phylloceras doderleinianum (CATULLO) (208.2011), Kis-Gerecse, Bed 102, O. douvillei horizon, D: 86, H: appr. 54 (62.7%), W:
28 (51.8%), U: ?.
78 Calliphylloceras propinquum GCZY (209.2011), Kis-Gerecse, Bed 102, O. douvillei horizon, D: 138, H: 82 (59.4%), W: 36
(44%), U: appr. 7 (5%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 11
Plate II

Fragmenta Palaeontologica Hungarica 29, 2011

12 KOVCS, Z.
1970: Calliphylloceras nilssoni (HBERT) PANNUZI, p. 66, pl. 4, figs 112.
1975: Calliphylloceras nilssoni (HBERT) DEZI & RIDOLFI, p. 15, figs 2224.
1986: Calliphylloceras nilssoni (HBERT) GAKOVI, p. 113, pl. 18, fig. 1ab (cum syn.).
2000: Calliphylloceras nilssoni (HBERT) JOLY, p. 85, text-figs 177181, pl. 19, figs 69 (cum syn.).
2007: Calliphylloceras nilssoni (HBERT) TOMAS & PLFY, p. 243, fig. 4/ef.
2007: Calliphylloceras nilssoni (HBERT) RULLEAU, p. 50, pl. 3, figs 2, 3.

Material 7 moderately preserved internal moulds Remarks The specimens are close to the type
from the H. serpentinum Zone. (Ammonites calypso, DORBIGNY 1845, pl. 110, figs 13, the
Description Medium-size, involute, compressed lectotype was designated by JOLY 1994, p. 97, pl. 32, fig.
form with suboval whorl-section. The umbilicus is narrow, 3ac). C. nilssoni is remarkably similar to C. capitanii, however,
the flanks are slightly convex, the venter is rounded. No the latter bears much shallower and curved constrictions
whole body chamber is preserved. The ornamentation (see above). According to JOLY (2000: 85), C. aveyronense
consists of 56 pronounced, slightly curved constrictions (MENEGHINI) is a synonym of C. nilssoni. C. aveyronense
on the last whorl. They rise from the umbilicus with was figured by GCZY from Bakonycsernye (1967a, pl. 5,
rectiradiate direction on the lower dorso-lateral part, then figs 6, 7).
curve forward, become shallower on the ventrolateral Distribution Europe, North Africa, Turkey,
part, and pass over the venter. The specimens bear Caucasus, Iran, North America, Japan. Hungary: Mecsek
almost straight constrictions on the inner whorls. The Mts, Bakony Mts; Gerecse Mts: H. serpentinum H.
suture-line resembles that of C. capitanii. bifrons Zones: all studied Gerecse sections.

Calliphylloceras propinquum GCZY, 1967

(Plate II: 78, Plate V: 78)
? 1966: Calliphylloceras nilssoni mediojurassica (PRINZ) KOLLROVANDRUSOV, p. 35, text-figs 8 9, pl. 2, fig. 4.
v 1967a: Calliphylloceras mediojurassicum propinquum n. subsp. GCZY, p. 33, text-fig. 29, pl. 9, fig. 4, pl. 63, fig. 32.
v 1967a: Calliphylloceras aff. mediojurassicum (PRINZ) GCZY, p. 34, text-fig. 30, pl. 9, fig. 5.

Material 5 internal moulds of different state of and diphyllic LS26.

preservation. Remarks The specimens are close to the type
Description Large, involute, compressed form with (GCZY 1967a, pl. 9, fig. 4) in morphology; however, the
suboval whorl-section. The flanks are moderately convex, convex umbilical margin of the type is not characteristic.
the venter is broad, low and rounded. No whole body The species differs from C. mediojurassicum (PRINZ) in
chamber is preserved. The ornamentation is characterized wider section, and in curved, regularly and widely spaced
by about 6 slightly curved, prorsiradiate constrictions. They constrictions; as well as from C. supraliassicum (POMPECKJ) in
emerge from the umbilicus with radiate direction, curved well-developed, regularly spaced constrictions (see RULLEAU
forward on the lowermost part of the flank, become 1998a, pl. 4, fig. 6, pl. 5, figs 13). The species also differs
shallower from the mid-height, and almost disappear from C. capitanii in slightly less developed constrictions,
crossing the venter. Fine, dense, straight and rectiradiate which, on the other hand, are deeper than those of C. spadae.
riblets can be traced on well-preserved specimens. The Distribution Slovakia; Hungary: Bakony Mts,
suture-line consists of wide and short E, wide, long and Gerecse Mts: H. falciferum Subzone: Kis-Gerecse and
slightly asymmetrical L lobes, diphyllic ES, triphyllic LS1, Nagy-Pisznice Sections.

Calliphylloceras spadae (MENEGHINI, 1875)

(Plate III: 3, Plate IV: 12)
1875: Phylloceras Spadae MENEGHINI, p. 106.
18671881: Ammonites (Phylloceras) Spadae n. sp. MENEGHINI, p. 93, pl. 19, figs 14.
v 1967a: Calliphylloceras spadae (MENEGHINI) GCZY, p. 28, text-fig. 22, pl. 7, figs 23, pl. 63, figs 24, 25 (cum syn.).
1970: Calliphylloceras spadae (MENEGHINI) PANNUZI, p. 73, pl. 5, figs 46.
2002a: Calliphylloceras cf. spadae (MENEGHINI) MACCHIONI, p. 53, fig. 21.

Explanation to Plate III

12 Lytoceras francisci (OPPEL) (216.2011), Nagy-Pisznice, Bed 134, O. douvillei horizon, D: 86, H: 36 (41.8%), W: 24 (66.6%), U: 28
(32.5%).
3 Calliphylloceras spadae (MENEGHINI) (218.2011), Kis-Gerecse, Bed 109, H. pseudoserpentinum horizon, D: 90, H: appr. 54
(60%), W: 24 (44.4%), U: appr. 7 (7.7%).
4 Lytoceras francisci (OPPEL) (214.2011), Kis-Gerecse, Bed 102, O. douvillei horizon, D: 180, H: 75 (41.6%), W: 40 (53.3%), U: 60
(33.3%).
56 Calliphylloceras capitanii (CATULLO) (211.2011), Kis-Gerecse, Bed 100, H. sublevisoni horizon, D: 65, H: appr. 34 (52.3%), W:
18 (64.2%), U: ?.

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 13
Plate III

Fragmenta Palaeontologica Hungarica 29, 2011

14 KOVCS, Z.
Material 10 poorly preserved internal moulds. by MENEGHINI (18671881, pl. 19, figs 1, 2, 3, 4; the type
Description Medium-size, involute form with wide was designated by PINNA 1969, pl. 6, fig. 10), and with the
suboval to rounded subrectangular whorl-section. The specimens in the literature. They are also close to Calliphyllo-
umbilicus is narrow and deep, the flanks are moderately ceras spadae chiesai (NEGRI) (NEGRI 1936, p. 15, pl. 2, figs 2,
convex, the venter is broad, low and rounded. No whole 3; GCZY 1967a, p. 30, fig. 23, pl. 7, fig. 5, pl. 63, fig. 26).
body chamber is preserved. The ornamentation consists of The species resembles C. capitanii in morphology, but differs
89 very shallow, narrow and prorsiradiate lateral constric- in straighter and shallower constrictions. The constrictions
tions. The suture-line is characterised by wide E, wide and of C. selinoides (MENEGHINI) are denser and narrower.
long, slightly asymmetrical L lobes, diphyllic ES, triphyllic Distribution Italy, Austria, Greece. Hungary:
LS1, and diphyllic LS24. Bakony Mts; Gerecse Mts: H. pseudoserpentinum horizon:
Remarks The specimens agree with those figured Kis-Gerecse and Crocodile Sections.

Suborder Lytoceratina HYATT, 1889

Superfamily Lytoceratoidea NEUMAYR, 1875
Family Lytoceratidae NEUMAYR, 1875
Genus Lytoceras SUESS, 1865
Lytoceras francisci (OPPEL, 1865)
(Plate III: 12, 4)
1856: Ammonites fimbriatus HAUER, p. 62, pl. 22, figs 12.
1865: Ammonites Francisci OPPEL, p. 551.
1930: Lytoceras Francisci OPPEL MITZOPOULOS, pl. 3, fig. 1.
1963: Lytoceras francisci (OPPEL) ZANZUCCHI, p. 113, pl. 14, fig. 2.
1966: Thysanoceras francisci OPPEL NUTSUBIDZE, pl. 13, figs 12.
1968: Lytoceras francisci (OPPEL) PINNA, p. 10, pl. 1, figs 9, 16, 18, 19, pl. 1 n.t., fig. 1, pl. 2 n.t., fig. 4 (cum syn.).
1975: Lytoceras francisci (OPPEL) DEZI & RIDOLFI, p. 12, figs 79.
2001: Lytoceras francisci (OPPEL) VENTURI & FERRI, p. 90, figs fg.

Material from the H. serpentinum Zone 4 (HAUER 1856, pl. 22, figs 12, designated by OPPEL
specimens in different state of preservation. 1865, p. 551), and agree well with those figured in the
Description Large, evolute, compressed form with literature. The species differs from L. cornucopia (YOUNG et
elliptical whorl-section. The umbilicus is wide and shallow BIRD) by slower growth up to 35 mm diameter, by shallower
on the inner, while deeper on the external whorls. The umbilicus of the innermost whorls, and by absence of
flanks are convex, the venter is broad, low and rounded. constriction. L. siemensi (DENCKMANN) is also a similar,
The growing of the innermost whorls is slow and steady, but compressed form, but differs from L. francisci in steadily
becomes faster from 35 diameter. No whole body chamber growing, much higher inner whorls. The Aalenian L. sub-
is preserved. The whorls are smooth, constrictions are francisci STURANI (= L. vaceki GCZY) differs in wider
absent. The lytoceratid suture-line consists of short and umbilicus, and in wide-oval section.
narrow E, and long, ramified, asymmetric L lobes. Both the Distribution Italy, Austria, Greece, Albania,
ES and the LS are well-developed and ramified. Caucasus. Hungary: Bakony Mts; Gerecse Mts: Lower
Remarks The specimens are close to the type Upper Toarcian: all studied Gerecse sections.

Lytoceras cornucopia (YOUNG et BIRD, 1822)

(Plate IV: 34)
1822: Ammonites cornucopia YOUNG & BIRD, p. 252, pl. 12, fig. 8.
1968: Lytoceras cornucopia (YOUNG et BIRD) PINNA, p. 6, pl. 1, fig. 20, pl. 2 n.t., fig. 1 (cum syn.).
1994: Lytoceras cornucopia (YOUNG et BIRD) MOUTERDE & MATTEI, p. 88, pl. 30, fig. 2, pl. 31, fig. 1.
1998a: Lytoceras cornucopia (YOUNG et BIRD) RULLEAU, p. 39, text-figs 6/35, pl. 9, figs 13, pl. 10, figs 13, pl. 11, figs 13 (cum syn.).
2001: Lytoceras cornucopia (YOUNG et BIRD) VENTURI & FERRI, p. 89.
2007: Lytoceras cornucopia (YOUNG et BIRD) RULLEAU, p. 53, figs 11/35, pl. 4, figs 56, pl. 5, fig. 7, pl. 6, figs 12.
2010: Lytoceras cornucopia (YOUNG et BIRD) DELSATE & WEIS, pl. 4, fig. 2.

Explanation to Plate IV
12 Calliphylloceras spadae (MENEGHINI) (219.2011), Kis-Gerecse, Bed 106, H. pseudoserpentinum horizon, D: 96, H: 58 (60.4%),
W: 34 (58.6%), U: ?.
34 Lytoceras cornucopia (YOUNG & BIRD) (213.2011), Kis-Gerecse, Bed 112, H. striatus horizon, D: 140, H: 56 (40%), W: 36
(64.2%), U: 51 (36.4%).
5 Dactylioceras (Orthodactylites) aff. directum (BUCKMAN) (220.2011), Kis-Gerecse, Bed 113, H. striatus horizon, D: 75, H:
20 (26.6%), W: 18 (90%), U: ?.
67 Mesodactylites aff. annulatiformis (BONARELLI) (217.2011), Kis-Gerecse, Bed 112, H. striatus horizon, D: 78, H: 18 (23%),
W: 18, U: ?.

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 15
Plate IV

Fragmenta Palaeontologica Hungarica 29, 2011

16 KOVCS, Z.
Material from the H. serpentinum Zone 13 poorly MONESTIER 1931, pl. 6, figs 1, 3, 1216, 20, 22, 24, 26
preserved internal moulds. 28; GCZY 1967a, pl. 19, fig. 1; SCHLEGELMILCH 1976,
Description Large, evolute form with wide suboval pl. 3, fig. 4; and RULLEAU 1998a). The one figured here
whorl-section. The umbilicus is wide and deep, the flanks with wide and shallow constrictions is close to those
are convex, the venter is broad, low and rounded. The figured by MOUTERDE & MATTEI (1994, pl. 30, fig. 2),
growing of the whorls is slower and steadier than that of L. and RULLEAU (1998a, pl. 11, fig. 2). Taking into consi-
francisci. No body chamber is preserved. The ornamenta- deration the variability of the species, L. ktenasi MITZO-
tion consists of 5 wide, shallow, and concave constrictions POULOS (MITZOPOULOS 1930, p. 31, pl. 3, fig. 3) is
on the last whorl. The suture-line is characteristically lyto- regarded here as a Mediterranean morphotype of L.
ceratid, but cannot be traced in all details. cornucopia. L. ktenasi was treated and figured by GCZY
Remarks The specimens figured in the literature (1967a, pl. 18, fig. 2), and by PINNA (1968, pl. 1, fig. 17).
show high variability in whorl-section and presence of Distribution Europe, North Africa, Turkey,
constrictions (see the type: YOUNG & BIRD 1822, pl. 12, Caucasus. Hungary: Mecsek Mts, Bakony Mts; Gerecse
fig. 8, refigured by BUCKMAN 1923, pl. 391A; and the Mts: H. serpentinum H. bifrons Zones: all studied
specimens figured by PRINCIPI 1915, pl. 17, fig. 1; Gerecse sections.

Suborder Ammonitina HYATT, 1899

Superfamily Eoderoceratoidea SPATH, 1929
Family Dactylioceratidae HYATT, 1867
Subfamily Dactylioceratinae HYATT, 1867
Genus Dactylioceras HYATT, 1867
Dactylioceras (Orthodactylites) aff. directum (BUCKMAN, 1926)
(Plate IV: 5)
1926: Orthodactylites directus nov. BUCKMAN, pl. 654.
1927: Orthodactylites mitis nov. BUCKMAN, pl. 738.
1957: Dactylioceras directus BUCKMAN MAUBEUGE, p. 216, pl. 26, figs 5256.
1972: Dactylioceras (Orthodactylites) directum (BUCKMAN) SCHMIDTEFFING, p. 107, pl. 7, fig. 4, pl. 19, fig. 14.
1981: Dactylioceras (Orthodactylites) directum (BUCKMAN) HILLEBRANDT & SCHMIDTEFFING, p. 34, text-figs 13, 16/f, pl. 1, figs 811 (cum syn.).
2007: Dactylioceras (Orthodactylites) mitis BUCKMAN RULLEAU, p. 64, pl. 13, fig. 6.

Material 2 fragmentary specimens. (Orthodactylites) semicelatum (SIMPSON) by HOWARTH (1980),

Description Moderately evolute form with suboval but the latter differs in more involute coiling and wider
whorl-section. Both the umbilical wall and the venter are whorl. On the other hand, the ranges of the two species
rounded, the flanks are convex. The preserved part of the are almost identical, O. directum is known from the upper
body chamber is one whorl in length. The ornamentation Dactylioceras tenuicostatumHarpoceras serpentinum Zones,
consists of dense, narrow, sharp and rectiradiate ribs passing and O. semicelatum, although it is typical for the upper
the venter. Almost all second ribs bifurcate at about mid- Dactylioceras tenuicostatum Zone, has been also docu-
hight without tubercles. The suture-line cannot be traced. mented from the lower Harpoceras serpentinum Zone of
Remarks D. (Orthodactylites) directum seems to be the Spain (ELMI et al. 1989; GOY & MARTNEZ 1990). O. mitis
most closely allied form to the Gerecse specimens. They was recognized as synonym of O. directum by SCHMIDT
are similar to the type (BUCKMAN 1926, pl. 654, refigured EFFING (1972), HOWARTH (1973), and HILLEBRANDT &
by HOWARTH 1980, pl. 80, figs 34) and the specimens SCHMIDTEFFING (1981).
figured in the literature in morphology; the poor state of Distribution England, France, Spain, Chile, North
preservation, however, does not allow the certain arrange- America, Tibet. Gerecse Mts: H. striatus horizon: Kis-
ment. The species was considered as a synonym of D. Gerecse and Bnya-hegy B Sections.

Subfamily Nodicoeloceratinae VENTURI & FERRI, 2001

Genus Catacoeloceras BUCKMAN, 1923

Explanation to Plate V
12 Nodicoeloceras cf. choffati (RENZ) (222.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 46, H: 12 (26%), W: 30 (250%),
U: 19 (41.3%).
34 Catacoeloceras crassum (YOUNG & BIRD) (215.2011), Bnya-hegy B, Bed 1, H. lusitanicum horizon, D: 50, H: 14 (28%), W: 24
(170%), U: 24 (48%).
5 Fontanelliceras cf. fontanellense (GEMMELLARO) (242.2011), Kis-Gerecse, Bed 113a, D. tenuicostatum Zone, D: 34, H:
appr. 10 (29.4%), W: ?, U: appr. 16 (47%).
6 Canavaria sp. (257.2011), Bnya-hegy B, Bed 16, E. emaciatum Zone, (L: 45, H: appr. 12).
78 Calliphylloceras propinquum GCZY (212.2011), Nagy-Pisznice, Bed 137, H. pseudoserpentinum horizon, D: 200, H: 123 (61.5%),
W: 54 (44%), U: 7 (3.5%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 17
Plate V

Fragmenta Palaeontologica Hungarica 29, 2011

18 KOVCS, Z.
Catacoeloceras crassum (YOUNG & BIRD, 1828)
(Plate V: 34)
1828: Ammonites crassus YOUNG & BIRD, p. 253.
1966: Catacoeloceras crassum (YOUNG & BIRD) PINNA, p. 99, text-fig. 7, pl. 5, figs 10, 1921 (cum syn.).
1985: Catacoeloceras crassum (YOUNG & BIRD) HENGSBACH, p. 376, pl. 7, figs 2021.
1991: Catacoeloceras crassum (YOUNG & BIRD) JIMNEZ & RIVAS, p. 174, pl. 7, figs 811.
1998b: Catacoeloceras crassum (YOUNG & BIRD) RULLEAU, p. 5, pl. 7, figs 12, 1112.
2008: Catacoeloceras crassum (YOUNG & BIRD) METODIEV, p. 98, fig. 3/l, m.
2010: Catacoeloceras crassum (YOUNG & BIRD) DELSATE & WEIS, pl. 2, figs 1, 4.

Material from the Bnya-hegy B Section 1 moder- secondaries cross the venter. The suture-line is richly
ately preserved internal mould. ornate with long and ramified E and L lobes, and simpler,
Description Evolute, depressed form with oblong- straight U lobe.
elliptic whorl-section. The umbilicus is deep, both the Remarks The specimen figured here is very close
umbilical and the lateral walls are convex; the venter is to the type (PHILLIPS 1829, pl. 12, fig. 15, refigured by
low, broad and rounded. No body chamber is preserved. BUCKMAN 1918, pl. 119, and SCHLEGELMILCH 1976, pl.
The ornamentation consists of regular, pronounced and 40, fig. 4). A detailed analysis of genus Catacoeloceras was
radial ribbing with tubercles. Primary ribs emerge from lately accomplished by HENGSBACH (1985).
the umbilicus, and bifurcate from tubercles at the Distribution Europe, North Africa, North
shoulder. The intercosta is wider than the rib-width. The America, East Siberia. Gerecse Mts: H. bifrons Zone:
specimen bears 36 primaries on the last whorl. The Bnya-hegy AB Sections.

Genus Nodicoeloceras BUCKMAN, 1926

Nodicoeloceras tuberculatum (KOTTEK, 1963)
(Plate VI: 34)
1963: Catacoeloceras tuberculatum n. sp. KOTTEK, p. 135, text-fig. 65, pl. 17, figs 13.
1964: Peronoceras subarmatum (YOUNG & BIRD) STANKIEVITCH, p. 15, pl. 1, fig. 6.
v 1966: Peronoceras baconicum n. sp. GCZY, p. 438, pl. 1, fig. 2.
1971: Nodicoeloceras baconicum (GCZY) PINNA & LEVISETTI, p. 105, p. 130, text-figs 12/BC, 14/P, 21/14, pl. 6, figs 5, 7, 8, pl. 7, figs 46.
1997: Nodicoeloceras baconicum (GCZY) PETTINELLI et al., pl. 3, fig. 3.
2001: Nodicoeloceras baconicum (GCZY) VENTURI & FERRI, p. 145.
v 2007: Catacoeloceras tuberculatum KOTTEK GCZY & SZENTE, pl. 1, figs 45.

Material from the H. serpentinum Zone 11 considered as a synonym of Nodicoeloceras angelonii (RA-
internal moulds of different state of preservation. MACCIONI) by PINNA & LEVISETTI (1971); however, C.
Description Evolute form with depressed, rounded tuberculatum clearly differs by presence of the irregular
subrectangular whorl-section. The umbilicus is wide and fibulate ribbing. Based on the sculpture, P. baconicum
shallow, the umbilical wall is convex, the margin is rounded. GCZY was recognized as Nodicoeloceras by PINNA &
The flanks are convex, the shoulder is rounded, the LEVISETTI (l.c.); this arrangement is accepted here for
venter is broad and convex. The body chamber of the KOTTEKs taxon as well. The Gerecse specimens agree
figured specimen is more than one whorl in length. The well with the holotypes of the respective species (KOTTEK
ornamentation consists of well-developed, rectiradiate 1963, pl. 17, fig. 1; GCZY 1966, pl. 1, fig. 2) in size,
ribs passing the venter. Some ribs bifurcate from strong morphology, sculpture and range. The P. subarmatum figured
tubercles at the shoulder. On the last whorl one or two by STANKIEVITCH (1964, pl. 1, fig. 6) differs from the
simple ribs alternate with one bifurcating rib, and a few type (YOUNG & BIRD 1822, pl. 13, fig. 3, refigured by
fibulate primaries occur irregularly. The suture-line cannot HOWARTH 1962, pl. 17, figs 5ab) in less developed
be traced in details. nodes on the inner whorls and irregular fibulation, this
Remarks Comparing the descriptions and the specimen was correctly considered as N. tuberculatum by
figures of types of C. tuberculatum (KOTTEK 1963) and P. PINNA & LEVISETTI (l.c.).
baconicum (GCZY 1966), the identity of the two forms is Distribution Greece, Italy, Caucasus, North Africa.
undoubtful; therefore GCZYs species is to be regarded Hungary: Bakony Mts; Gerecse Mts: O. douvillei horizon
as a junior synonym of C. tuberculatum. The latter was H. sublevisoni Subzone: all studied Gerecse sections.

Explanation to Plate VI
12 Mesodactylites mediterraneus (RENZ) (225.2011), Bnya-hegy B, Bed 4, H. pseudoserpentinum horizon, D: 76, H: 21 (27.6%), W:
26 (123%), U: 40 (52.6%).
34 Nodicoeloceras tuberculatum (KOTTEK) (221.2011), Nagy-Pisznice, Bed 133, O. douvillei horizon, D: 92, H: 21 (22.8%), W: 23
(109.5%), U: 50 (54.3%).
5 Harpoceras (Harpoceras) serpentinum (SCHLOTHEIM) (223.2011), Bnyahegy B, Bed 12, H. striatus horizon, D: 180, H:
66 (36.6%), W: 22 (33.3%), U: 60 (33.3%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 19
Plate VI

Fragmenta Palaeontologica Hungarica 29, 2011

20 KOVCS, Z.
Nodicoeloceras cf. choffati (RENZ, 1912)
(Plate V: 12)
1912: Coeloceras Choffati n. sp. RENZ, p. 86, pl. 6, fig. 5.
1971: Nodicoeloceras choffati (RENZ) PINNA & LEVISETTI, p. 100, p. 128, text-figs 14/M, 18/A, 21/26, pl. 4, figs 37, 10, 13 (cum syn.).
1972: Coeloceras choffati RENZ SCHMIDTEFFING, p. 83, pl. 1, fig. 2.
1973: Nodicoeloceras (?) choffati (RENZ) GUEX, p. 510, pl. 12, fig. 4.
1975: Nodicoeloceras choffati (RENZ) DEZI & RIDOLFI, p. 40, fig. 108.
1980: Nodicoeloceras choffati (RENZ) WIEDENMAYER, p. 81, pl. 31, fig. 4.
1981: Peronoceras cf. choffati (RENZ) HILLEBRANDT & SCHMIDTEFFING, p. 51, pl. 4, figs 56.
1998: Nodicoeloceras aff. choffati (RENZ) KMENT, p. 185, text-fig. 8, pl. 2, fig. 1.

Material 1 poorly preserved internal mould. width. The secondaries bend slightly forward, and cross
Description Evolute form with deep umbilicus. Both the venter. The suture-line is intricate, but cannot be
the umbilical and the lateral walls are oblique and slightly traced in all details.
convex. The ventrolateral shoulder is well-pronounced; the Remarks The specimen is similar to the type (RENZ
venter is low, broad and rounded. The whorl-section is 1912, pl. 6, fig. 5) and to those figured in the literature, but
wide, rounded diamond-shaped, it agrees well with that the poor state of preservation does not allow the certain
of the specimen figured by KMENT (1998). No body identification.
chamber is preserved. Radial primary ribs rise from the Distribution Portugal, France, Italy, Greece, Austria,
umbilicus, and bi- or trifurcate from pronounced tubercles North Africa, South America. Gerecse Mts: H. striatus
at the shoulder. The intercosta is as wide as the rib- horizon: Bnya-hegy B Section.

Genus Mesodactylites PINNA & LEVISETTI, 1971

Mesodactylites mediterraneus (RENZ, 1912)
(Plate VI: 12)
1912: Coeloceras Desplacei DORB. n. var. mediterranea RENZ, p. 68.
1971: Mesodactylites mediterraneus (MEISTER) PINNA & LEVISETTI, p. 93, p. 127, text-fig. 21/1, pl. 2, figs 9, 13, 14, pl. 3, fig. 9 (cum syn.).
1975: Mesodactylites mediterraneus (MEISTER) DEZI & RIDOLFI, p. 39, fig. 105.
2001: Mesodactylites mediterraneus (RENZ) VENTURI & FERRI, p. 153, figs gi.

Material 1 moderately preserved internal mould. venter. The suture-line cannot be traced in all details.
Description Medium-size, evolute form with shallow Remarks The specimen agrees with the type
umbilicus. Both the umbilical margin and the ventrolateral (figured by MEISTER 1913, pl. 15, fig. 1). M. annulatiformis
shoulder are rounded, the flanks are convex. The venter is (BONARELLI) is a similar form in morphology (KOTTEK
broad and low, the whorl-section is depressed wide oval. 1963, p. 128), but differs in more pronounced ribbing.
The preserved body chamber is one whorl in length. The Nodicoeloceras crassoides (SIMPSON) differs in wider whorls.
ornamentation consists of moderately developed, radial Distribution Portugal, Spain, Italy, Greece, Algeria.
ribs without any nodes. Simple and bifurcating ribs Gerecse Mts: uppermost H. pseudoserpentinum horizon:
alternate with each other almost regularly; and cross the Bnya-hegy B Section.

Mesodactylites aff. annulatiformis (BONARELLI, 1899)

(Plate IV: 67)
18671881: Ammonites (Stephanoceras) Desplacei DORBIGNY MENEGHINI, p. 76, pl. 16, figs 78.
1899: Coeloceras annulatiforme BONARELLI, p. 212.
1943: Coeloceras (Dactylioceras) annulatiforme BONARELLI MAXIA, p. 113, pl. 3, fig. 4.
1971: Mesodactylites annulatiformis (BONARELLI) PINNA & LEVISETTI, p. 93, p. 127, text-figs 14/G, 21/5, pl. 2, fig. 10 (cum syn.).
1972: Nodicoeloceras annulatiforme (BONARELLI) SCHMIDTEFFING, p. 64, text-fig. 12.
2004: Nodicoeloceras ? annulatiforme (BONARELLI) MORARD, p. 267.

Material 3 poorly preserved internal moulds. characterised by dense, sharp and rectiradiate ribs
Description Evolute form with subcircular passing the venter. Some primaries bifurcate without
whorl-section. The umbilicus is wide and shallow, the any tubercles above the mid-height. 12 simple ribs
margin is rounded, the flanks are convex, the venter is occur between the bifurcating ribs. The suture-line
broad, low and rounded. The ornamentation is cannot be traced.

Explanation to Plate VII

1 Praepolyplectus sp. (227.2011), Bnya-hegy B, Bed 13, H. striatus horizon, D: 74, H: 40 (54%), W: 12 (30%), U: ?.
2 Harpoceras (Harpoceras) falciferum (SOWERBY) (224.2011), Kis-Gerecse, Bed 110, H. pseudoserpentinum horizon, L: 104, H: ?, W: 16, U: ?.
3 Harpoceras (Harpoceras) pseudoserpentinum GABILLY (226.2011), Nagy-Pisznice, Bed 141, H. pseudoserpentinum horizon, D: 200, H:
60 (30%), W: 26 (43.3%), U: 95 (47.5%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 21
Plate VII

Fragmenta Palaeontologica Hungarica 29, 2011

22 KOVCS, Z.
Remarks The specimens are close to the lecto- ites) semiannulatus HOWARTH is a closely allied form, but
type (MENEGHINI 18671881, pl. 16, fig. 7, refigured by differs in wider whorls.
PINNA 1969, pl. 4, figs 23), and those figured in the Distribution Italy, Portugal, Greece, North Africa,
literature; however, due to the poor preservation, none (?)Austria, (?)northern Spain. Gerecse Mts: H. striatus
of them can be arranged with certainty. D. (Orthodactyl- horizon: Kis-Gerecse and Bnya-hegy B Sections.

Superfamily Hildoceratoidea HYATT, 1867

Family Hildoceratidae HYATT, 1867
Subfamily Arieticeratinae HOWARTH, 1955
Genus Fontanelliceras FUCINI, 1931
Fontanelliceras cf. fontanellense (GEMMELLARO, 1886)
(Plate V: 5)
1886: Harpoceras Fontanellense GEMMELLARO, p. 118, pl. 2, figs 12.
2003: Fontanelliceras fontanellense (GEMMELLARO) MACCHIONI & MEISTER, p. 398, pl. 10, figs 1, 4 (cum syn.).
2004: Fontanelliceras fontanellense (GEMMELLARO) MORARD, p. 287, pl. 9, figs 1, 6, 25.
2010: Fontanelliceras fontanellense (GEMMELLARO) BILOTTA et al., fig. 12/H.

Material 1 poorly preserved internal mould. 146, fig. 88/a); however, the state of preservation does
Description Small, evolute, platycone form with not allow the exact arrangement. Although the species is
moderately deep umbilicus. Both the margin and the characterised by a long stratigraphic range, the lithofacies
shoulder are rounded, the flanks are slightly convex. The of the figured specimen indicates the Dactylioceras
ribs are simple, strong and slightly sigmoid. The suture- tenuicostatum Zone of the Kis-Gerecse Section (see
line cannot be traced. above).
Remarks The Gerecse specimen resembles the Distribution The species is known from the E.
lectotype (GEMMELLARO 1886, pl. 2, fig. 1, refigured by emaciatumlower D. tenuicostatum Zones of Italy,
FUCINI 1931, pl. 8, fig. 21, and MACCHIONI 2002a, p. France, Spain, North Africa, eastern Russia, Japan.

Subfamily Polyplectinae VENTURI, 1981

Genus Praepolyplectus VENTURI, 1981
Praepolyplectus sp.
(Plate VII: 1)
Material 3 poorly preserved internal moulds. fig. 2; JAKOBS 1997, pl. 4, fig. 14; MYCZYNSKI 2004, fig.
Description Oxycone form with acutely sharpened 24/3; BCAUD 2006, pl. 4, fig. 2), as well as the range of
venter and narrow umbilicus. The flanks are slightly the species is typical of the MiddleUpper Toarcian
convex; the section is subtriangular. No whole body (NAY 2011). (The early occurrence of P. discoides in the
chamber is preserved. The simple, prorsiradiate, falcoid Serpentinum Zone [JIMNEZ & RIVAS 1992] needs more
ribs are wide, low and flat. The suture-line seems slightly research.) The Gerecse specimens are similar to the type
less intricate than that of Polyplectus, however, it cannot be of genus Praepolyplectus (P. forzanensis VENTURI, 1981, fig.
traced in all details. 7/2ab) in morphology. They are also close to the Prae-
Remarks The specimens differ from both Poly- polyplectus specimens figured by VENTURI (1982, p. 69),
plectus pluricostatus (HAAS) and P. apenninicus (HAAS) by FARAONI et al. (1994, pl. 13, fig. 14), PARISI et al. (1998,
possessing wider and flattened ribs. On the other hand, text-fig. 7/17, pl. 3, fig. 15), VENTURI & FERRI (2001, p.
the ribs of P. discoides (ZIETEN) are much wider (see the 189), and VENTURI et al. (2010, p. 278, 310); however,
type: ZIETEN 1831, pl. 16, fig. 1, refigured by HOWARTH due to the poor preservation, none of them can be
1992, pl. 28, fig. 7; and e.g. the specimens of DORBIGNY arranged with certainty at species level.
1846, pl. 115, fig. 1, refigured by MOUTERDE & GABILLY Distribution Italy, Greece, (?)southern Spain.
1994, pl. 35, fig. 3, and DUBAR & MOUTERDE 1965, pl. 1, Gerecse Mts: H. striatus horizon: Bnya-hegy B Section.

Explanation to Plate VIII

1 Polyplectus pluricostatus (HAAS) (230.2011), Kis-Gerecse, Bed 103, H. pseudoserpentinum horizon, D: 90, H: 50 (55.5%), W: 16
(32%), U: 9 (10%).
2 Harpoceras (Harpoceras) aff. mediterraneum PINNA (231.2011), Bnya-hegy B, Bed 9, H. pseudoserpentinum horizon, D: 112,
H: 42 (37.5%), W: 20 (47.6%), U: ?.
34 Harpoceras (Harpoceras) subplanatum (OPPEL) (237.2011), Kis-Gerecse, Bed 103, H. pseudoserpentinum horizon, D: 170, H:
80 (47%), W: 26 (32.5%), U: 33 (19.4%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 23
Plate VIII

Fragmenta Palaeontologica Hungarica 29, 2011

24 KOVCS, Z.
Genus Polyplectus BUCKMAN, 1890
Polyplectus pluricostatus (HAAS, 1913)
(Plate VIII: 1)
1906: Polyplectus n. f. aff. discoides ZIETEN PARISCH & VIALE, p. 150, pl. 8, fig. 4.
1913: Harpoceras (Polyplectus) discoides (ZIETEN) var. pluricostata HAAS, p. 117, pl. 6, fig. 3ab.
1965: Polyplectus pluricostatus (HAAS) DUBAR & MOUTERDE, p. 82, text-figs 1315, pls 23.
1969: Polyplectus discoides pluricostatus HAAS GALLITELLI WENDT, p. 22, pl. 7, fig. 7.
2001: Polyplectus pluricostatus HAAS VENTURI & FERRI, p. 190.
2006: Polyplectus pluricostatum (HAAS) BCAUD, p. 54, text-fig. 15B, pl. 5, fig. 2 (cum syn.).
v 2007: Polyplectus pluricostatus (HAAS) GALCZ et al., p. 345, fig. 3/45.
v 2008: Polyplectus pluricostatus (HAAS) GCZY et al., pl. 1, fig. 14.

Material from the H. serpentinum Zone 3 poorly Remarks The specimens agree well with the type
preserved internal moulds. (PARISCH & VIALE 1906, pl. 8, fig. 4), and with the speci-
Description Oxycone form with acutely angled mens figured in the literature. P. discoides bears wider ribs,
venter and very narrow umbilicus. The flanks are slightly and ranges in the Middle to Upper Toarcian (see above).
convex, almost flat; the section is subtriangular. The Distribution Europe, North Africa, South and
specimen figured here bears a well-developed keel. No (?)North America. Hungary: Bakony Mts; Gerecse Mts: H.
whole body chamber is preserved. The ribs are fine, striatus horizon H. sublevisoni Subzone: Bnya-hegy B
narrow and flexuous. The suture-line is richly ornate with Section, H. pseudoserpentinum horizon H. bifrons Zone:
four or five auxiliary saddles. Kis-Gerecse, Nagy-Pisznice and Crocodile Sections.

Subfamily Harpoceratinae NEUMAYR, 1875

Genus Harpoceras WAAGEN, 1869
Harpoceras (Harpoceras) serpentinum (SCHLOTHEIM, 1813)
(Plate VI: 5, Plate IX: 12, 8)
1813: Ammonites serpentinus SCHLOTHEIM, p. 35.
1818: Argonauta serpentinus REINECKE, p. 89, pl. 13, figs 7475.
1843: Ammonites alternatus SIMPSON, p. 43.
1992: Harpoceras serpentinum (SCHLOTHEIM) HOWARTH, p. 109, text-figs 18E, 2327, pl. 15, figs 3, 5, pl. 16, figs 15, pl. 17, figs 17, pl.
18, figs 12, pl. 19, fig. 1 (cum syn.).
2006: Harpoceras (Harpoceras) serpentinum (SCHLOTHEIM) BCAUD, p. 58, pl. 10, fig. 1, pl. 18, fig. 3 (cum syn.).
2007: Harpoceras serpentinum (SCHLOTHEIM) RULLEAU, p. 68, pl. 16, fig. 10, pl. 17, fig. 5.
2008: Harpoceras serpentinum (SCHLOTHEIM) METODIEV, fig. 4/d, e.

Material 9 internal moulds of different state of suture-line is characterised by narrow E, narrow and long L,
preservation. and small U lobes. The ES is wide and asymmetrically
Description Moderately evolute, compressed form divided, both the LS1 and LS2 are narrow and long.
with narrow elliptical to rounded subrectangular whorl- Remarks The species is one of the index fossils of
section. The umbilicus is wide and shallow, the margin is the Harpoceras serpentinum Zone, so its occurrance in
rounded. The flanks are slightly convex. A shallow, disconti- three Gerecse sections made it possible to establish the
nuous lateral groove appears at mid-height on some speci- Submediterranean ammonite stratigraphy for this Mediter-
mens, but it differs from the marked groove of H. falciferum ranean fauna. The lectotype was designated and figured by
(SOWERBY). The venter is narrow, rounded, and carinate HOWARTH (1992, p. 110, text-fig. 23), as well as a detailed
with a high keel. No whole body chamber is preserved. The history of nomenclature was accomplished (l.c., p. 114115).
ornamentation consists of well-developed falcoid ribs. The Summarily, SCHLOTHEIMs species has priority over that of

Explanation to Plate IX
12 Harpoceras (Harpoceras) serpentinum (SCHLOTHEIM) (235.2011) Bnya-hegy B, Bed 12, H. striatus horizon, D: 82, H: 34
(41.4%), W: 17 (50%), U: 24 (29.2%).
3 Ovaticeras cf. ovatum (YOUNG et BIRD) (233.2011) Bnya-hegy B, Bed 3, H. pseudoserpentinum horizon, D: 95, H: 36 (37.8%),
W: 20 (55.5%), U: 31 (32.6%).
4 Harpoceras (Harpoceras) mediterraneum PINNA (232.2011) Bnya-hegy B, Bed 3, O. douvillei horizon, D: 80, H: 32 (40%),
W: 18 (56.2%), U: 29 (36.2%).
5 Harpoceras (Harpoceras) subexaratum BONARELLI (228.2011) Bnya-hegy A, Bed 46, O. douvillei horizon, D: 70, H: 28
(40%), W: 12 (42.8%), U: 23 (32.8%).
67 Harpoceras (Harpoceras) strangewaysi (SOWERBY) (229.2011) Kis-Gerecse, Bed 112, H. striatus horizon, L: 128, H: 48, W:
24 (50%), U: ?.
8 Harpoceras (Harpoceras) serpentinum (SCHLOTHEIM) (236.2011) Bnya-hegy B, Bed 11, H. striatus horizon, D: 92, H: 36
(39%), W: 18 (50%), U: 29 (31.5%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 25
Plate IX

Fragmenta Palaeontologica Hungarica 29, 2011

26 KOVCS, Z.
REINECKE. The latter is to be considered as Harpoceras the specimen figured on Plate IX: 12 bears less falcoid ribs,
serpentinum, while most specimens described later in the it is close to that figured by HOWARTH (l.c., pl. 17, fig. 2).
literature under the name A. serpentinus REINECKE belong to Two specimens correspond to the type of A. alternatus
genus Hildaites (for synonyms see BCAUD 2006: 146). figured by BUCKMAN (1909, pl. 9, refigured by HOWARTH
According to HOWARTH (l.c.), Harpoceratoides alternatus l.c., pl. 16, fig. 1), the one figured on Plate IX: 8, and a
(SIMPSON) is a synonym of H. serpentinum; BCAUD (2006), fragment from Bed 110 of the Kis-Gerecse Section. Both
however, considered it as a subspecies of the latter. The also resemble the specimens of HOWART (l.c., pl. 18, fig. 1),
specimens figured in the literature show moderate variability and of RULLEAU (2007, pl. 17, fig. 5).
in shape and development of ribbing. The large Gerecse Distribution Europe, North Africa, Caucasus,
specimen (Plate VI: 5) agrees with the examples of Madagascar, North and South America. Hungary: Mecsek
HOWARTH (1992, text-fig. 26, pl. 16, fig. 2), and with one of Mts, Bakony Mts; Gerecse Mts: H. striatus horizon: Kis-
the transitional forms of KMENT (1998, pl. 2, fig. 2), while Gerecse, Nagy-Pisznice and Bnya-hegy B Sections.

Harpoceras (Harpoceras) strangewaysi (SOWERBY, 1820)

(Plate IX: 67)
1820: Ammonites strangewaysi SOWERBY, p. 99, pl. 254, figs 1, 3.
1976: Harpoceratoides strangewaysi (SOWERBY) GABILLY, p. 85, pl. 5, figs 12.
2006: Harpoceras (Harpoceras) strangewaysi (SOWERBY) BCAUD, p. 57, pl. 6, fig. 3, pl. 7, fig. 2, pl. 9, figs 1, 2 (cum syn.).

Material 1 fragmentary internal mould. ES is slightly asymmetric.

Description Moderately evolute, compressed form Remarks The taxon had been regarded as a
with rounded subrectangular whorl-section. The umbilicus synonym of Harpoceras serpentinum (SCHLOTHEIM) by
is deep, the umbilical wall is concave, the margin is rounded. HOWARTH (1992), but its validity was confirmed again by
The flanks are almost flat and parallel with a shallow, BCAUD (2002, 2006). The Gerecse specimen agrees well
discontinuous lateral groove under the mid-flank. The with the type (SOWERBY 1820, pl. 254, figs 1, 3, refigured
shoulder is rounded, the venter is wide, low, rounded and by HOWARTH 1992, pl. 15, figs 4ab), and with the
carinate. The ribbing is weakly developed, fine, dense and specimens figured in the literature. Despite of being a
falcoid, almost disappears on the dorsal half of the last fragment, it can be arranged with certainty.
whorl. The suture-line consists of narrow E, somewhat Distribution England, France, Germany, North
wide and long L, and short, narrow U lobes. The divided Africa. Gerecse Mts: H. striatus horizon: Kis-Gerecse Section.

Harpoceras (Harpoceras) pseudoserpentinum GABILLY, 1976

(Plate VII: 3)
1976: Harpoceras pseudoserpentinum GABILLY, p. 90, pl. 7, figs 1, 45, pl. 8, fig. 1, pl. 9, figs 12 (cum syn.) (non: pl. 6, figs 12 = H.
[Harpoceras] rulleaui BCAUD).
? 1990: Harpoceras pseudoserpentinum GABILLY GOY & MARTNEZ, pl. 1, fig. 7.
1994: Harpoceras pseudoserpentinum GABILLY GABILLY & MOUTERDE, p. 56, pl. 28, figs 3ab.
2006: Harpoceras (Harpoceras) pseudoserpentinum GABILLY BCAUD, p. 60, pl. 16, fig. 1, pl. 17, fig. 1 (cum syn.).
2007: Harpoceras pseudoserpentinum GABILLY RULLEAU, p. 68, pl. 17, fig. 1.
non 2010: Harpoceras pseudoserpentinum GABILLY GURINFRANIATTE et al., pl. 2, fig. 6.

Material 1 moderately preserved internal mould. remarkable in a Mediterranean fauna. The specimen figured
Description Large, evolute, compressed form. here agrees well with the type (GABILLY 1976, pl. 7, fig. 1,
The umbilicus is shallow, the slope is concave, the margin pl. 9, fig. 1), and with those of GABILLY & MOUTERDE
is rounded. The flanks are slightly convex with a wide, (1994), BCAUD (2006), and RULLEAU (2007). As a specific
shallow, discontinuous lateral groove close to the mid- morphological feature, three parts of almost equal size can
height. The carinate venter is wide, low, and rounded; the be distinguished on the flank: a slightly aslope and flattened
whorl-section is suboval. No whole body chamber is umbilico-lateral part; a relatively wide, discontinuous lateral
preserved. The ribbing is fine, moderately dense and groove that runs along the mid-flank, and a slightly convex
falcoid. The suture-line is characterized by narrow and ventro-lateral part. H. falciferum differs in more convex
short E, wide and long L, and moderately wide, short U flanks, narrower and continuous lateral groove.
lobes. The ES is broad and asymmetrically divided. Distribution France, Switzerland, (?)northern Spain,
Remarks The species is an important horizon (?)England. Gerecse Mts: H. pseudoserpentinum horizon: Nagy-
marker in the Submediterranean regions; its occurrence is Pisznice Section.

Explanation to Plate X
12 Harpoceras (Harpoceras) subexaratum BONARELLI (238.2011), Bnya-hegy B, Bed 5, H. pseudoserpentinum horizon, D: 122,
H: 50 (41%), W: 28 (56%), U: 37 (30.3%).
35 Harpoceras (Harpoceras) mediterraneum PINNA (234.2011), Kis-Gerecse, Bed 105, H. pseudoserpentinum horizon, L: 135, H:
50, W: 30 (60%), U: ?.

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 27
Plate X

Fragmenta Palaeontologica Hungarica 29, 2011

28 KOVCS, Z.
Harpoceras (Harpoceras) falciferum (SOWERBY, 1820)
(Plate VII: 2, Plate XI: 1)
1820: Ammonites falcifer SOWERBY, p. 99, pl. 254, fig. 2.
1964: Harpoceras serpentinum (REINECKE) STANKIEVITCH, p. 22, pl. 3, figs 12.
2006: Harpoceras (Harpoceras) falciferum (SOWERBY) BCAUD, p. 61, pl. 19, fig. 1, pl. 20, fig. 1, pl. 24, fig. 2, pl. 26, fig. 1, pl. 27, figs 1, 2, pl.
32, figs 2, 4 (cum syn.)
2006: Harpoceras falcifer (SOWERBY) TOPCHISHVILI et al., pl. 13, figs 13.
2006: Harpoceras mulgravium (YOUNG & BIRD) TOPCHISHVILI et al., pl. 14, fig. 1.
2008: Harpoceras falciferum (SOWERBY) METODIEV, fig. 4/f, g, h.

Material 3 moderately preserved internal moulds. 6, pl. 17, figs 16). The fragmentary specimen figured on
Description Large, moderately evolute, compressed Plate VII: 2 is close to the type (SOWERBY 1820, pl. 254,
form with narrow oval whorl-section. The umbilicus is fig. 2, refigured by e.g. SCHLEGELMILCH 1976, pl. 45, fig.
wide and shallow, the umbilical wall is flat and oblique, 4, and HOWARTH 1992, pl. 19, fig. 2). The other on Plate
the margin is rounded. The flanks are slightly convex XI: 1 bears denser and finer ribs than the type, it is close to
with a relatively wide, very shallow lateral groove under the type of H. concinnum (BUCKMAN 1927, pl. 742), and
the mid-height. The venter is narrow, rounded and carinate. resembles the specimen figured by RULLEAU (2007, pl. 18,
No whole body chamber is preserved. The ribbing is fig. 1). The typical form of the nominal species is known
dense and falcate. The suture-line consists of broad, from Mediterranean localities as well (e.g. Italy: NICOSIA &
slightly asymmetrically divided ES, broad, long L and PALLINI 1977; Austria: KMENT 1998; Caucasus: STAN-
short, straight U lobes. KIEVITCH 1964, KAZAKOVA 1987, TOPCHISHVILI et al.
Remarks According to HOWARTH (1992) and 2006), while H. concinnum was recorded from e.g. Italy
BCAUD (2006), Harpoceras mulgravium (YOUNG & BIRD) (ZANZUCCHI 1963; PINNA 1968), Greece (KOTTEK 1963),
and H. concinnum BUCKMAN are synonyms of H. falciferum. Morocco (GUEX 1973), and southern Spain. The H. n. sp.
The species is characterized by significant variability in the ? gr. falcifer figured by ZANZUCCHI (1963, pl. 18, fig. 10)
style of coiling and ribbing (see GABILLY 1976, pl. 5, figs corresponds with the morphology of H. mediterraneum, and
34, pl. 9, figs 35, pl. 11, figs 13, pl. 12, figs 12, pl. 14, probably represents a transitional form.
figs 12, pl. 15, pl. 16, figs 12; HOWARTH 1992, pl. 19, Distribution World-wide. Hungary: Mecsek Mts,
figs 24, pl. 20, figs 111, text-figs 3033; BCAUD 2002, Bakony Mts; Gerecse Mts: H. pseudoserpentinum horizon:
pl. 5, figs 13, 2006, l.c.; MORARD 2004, pl. 16, figs 1, 34, Kis-Gerecse and Nagy-Pisznice Sections.

Harpoceras (Harpoceras) mediterraneum PINNA, 1968

(Plate IX: 4, Plate X: 35)
18671881: Ammonites falcifer SOW. MENEGHINI, p. 14, pl. 3, figs 2ac.
1968: Harpoceras (Harpoceras) falcifer (SOWERBY) mediterraneum n. subsp. PINNA, p. 37, pl. 2, fig. 10, pl. 3, figs 5, 7, 8, 10, pl. 4, fig. 7, pl. 5,
fig. 1, pl. 6, figs 2, 3, pl. 1 n.t., figs 7, 12, 14, pl. 2 n.t., fig. 28 (cum syn.).
1969: Harpoceras subexaratum BONARELLI GALLITELLI WENDT, p. 22, pl. 7, fig. 4.
1973: Harpoceras mediterraneum (PINNA) GUEX, p. 500, pl. 2, fig. 2, pl. 4, fig. 5, pl. 14, fig. 27.
1975: Harpoceras falcifer (SOWERBY) mediterraneum (PINNA) DEZI & RIDOLFI, p. 17, figs 3133.
1979: Harpoceras falcifer (SOWERBY) mediterraneum PINNA MARIOTTI et al., pl. 1, fig. 8.
1992: Harpoceras (Harpoceras) mediterraneum PINNA JIMNEZ & RIVAS, p. 46, pl. 3, figs 35 (non: pl. 2, figs 12).
1994: Harpoceras mediterraneum PINNA FARAONI et al., pl. 13, figs 3, 15, pl. 17, fig. a.
1997: Harpoceras mediterraneum PINNA PETTINELLI et al., pl. 3, fig. 2.
2001: Harpoceras mediterraneum PINNA VENTURI & FERRI, p. 181, p. 187, figs ad.
v 2007: Harpoceras mediterraneum PINNA GALCZ et al., p. 345, fig. 3/1.
v 2008: Harpoceras mediterraneum PINNA GCZY et al., pl. 1, fig. 13.
2010: Harpoceras cf. mediterraneum PINNA VENTURI et al., p. 276, 303.

Material from the H. serpentinum Zone 7 internal somewhat broad and low, flattened on the body chamber,
moulds of different state of preservation. and carinate with a narrow, high keel. The whorl-section
Description Moderately involute, compressed form. is subovalsubrectangular with maximum thickness at the
The umbilicus is slightly deep, the umbilical wall is lower third. The ornamentation consists of strong and
undercut, the flanks are slightly convex, and both the falcoid ribs on the inner whorls that becomes weaker on
margin and the shoulder are rounded. The venter is the last whorl of the phragmocone and the body chamber.

Explanation to Plate XI
1 Harpoceras (Harpoceras) falciferum (SOWERBY) (240.2011), Nagy-Pisznice, Bed 140, H. pseudoserpentinum horizon, D:
108, H: 38 (35%), W: appr. 16 (42%), U: 39 (36%).
2 Hildaites striatus GUEX (241.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 98, H: 26 (26.5%), W: 16 (61.5%), U: 46
(47%).
3 Harpoceras (Harpoceras) subplanatum (OPPEL) (239.2011), Nagy-Pisznice, Bed 134, O. douvillei horizon, D: 170, H: 80
(47%), W: 28 (35%), U: 34 (20%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 29
Plate XI

Fragmenta Palaeontologica Hungarica 29, 2011

30 KOVCS, Z.
No whole body chamber is preserved. The suture-line is subexaratum. Indeed, the morphology of H. mediterraneum
characterized by somewhat wide E, and narrow and long seems to show affinity with the H. serpentinum specimen
L lobes, wide, divided and slightly asymmetrical ES, and figured here on Plate IX: 12; while some H. mediterra-
narrower L2. neum specimens bearing less developed ribs (e.g. GUEX
Remarks The Gerecse specimens are very close to 1973, pl. 4, fig. 5; PETTINELLI et al. 1997, pl. 3, fig. 2)
the holotype (PINNA 1968, pl. 2, fig. 10, refigured by may be related to H. subexaratum. The H. subexaratum
PINNA 1969, pl. 1, fig. 12, and PINNA & SPEZIA 1975, pl. presented by GALLITELLI WENDT (1969) agrees well
12, fig. 4). The species is a characteristic Mediterranean with the holotype of H. mediterraneum, it is regarded here
counterpart of H. falciferum; both appeared in the lower as an example of the latter taxon. H. mediterraneum is used
H. pseudoserpentinum horizon. H. mediterraneum differs from here as a subzone marker for the Bnya-hegy B Section.
H. falciferum in more rounded ventral part, absence of Distribution Italy, southern Spain, Greece, Albania,
lateral groove, and bearing slightly finer, less falcoid ribs. North Africa. Hungary: Bakony Mts; Gerecse Mts: H.
According to GABILLY (1976), H. mediterraneum derived falciferum Subzone H. bifrons Zone: all studied
from the H. serpentinum group, and was the ancestor of H. Gerecse sections.

Harpoceras (Harpoceras) aff. mediterraneum PINNA, 1968

(Plate VIII: 2)
Material 2 fragmentary specimens. ribbing resembles that of the specimen figured by GUEX
Remarks Both are closely allied in morphology (1973, pl. 5, fig. 5).
with Harpoceras (Harpoceras) mediterraneum, but differ in Distribution H. pseudoserpentinum horizon: Bnya-
wider and somewhat more convex ribs. This style of hegy B and Crocodile Sections.

Harpoceras (Harpoceras) subexaratum BONARELLI, 1899

(Plate IX: 5, Plate X: 12)
18671881: Ammonites complanatus BRUG. MENEGHINI, p. 16, pl. 4, figs 1ac.
1899: Harpoceras subexaratum n. f. BONARELLI, p. 201.
v 1967b: Polyplectus cf. subexaratus (BONARELLI) GCZY, p. 122, text-fig. 5, pl. 1, fig. 6, pl. 2, fig. 2.
1968: Harpoceras (Harpoceras) exaratum (YOUNG & BIRD) subexaratum BONARELLI PINNA, p. 40, pl. 4, fig. 2, pl. 5, figs 2, 5, 10, pl. 6, fig. 1,
pl. 1 n.t., figs 4, 13, 16, 19, pl. 2 n.t., fig. 29 (cum syn.).
? 1968: Harpoceras (Harpoceras) cf. mulgravium (YOUNG & BIRD) PINNA, p. 39, pl. 5, fig. 9, pl. 1 n.t., fig. 15, pl. 2 n.t., fig. 33.
non 1969: Harpoceras subexaratum BONARELLI GALLITELLI WENDT, p. 22, pl. 7, fig. 4 (= H. mediterraneum PINNA).
1975: Harpoceras exaratum (YOUNG & BIRD) subexaratum (BONARELLI) DEZI & RIDOLFI, p. 17, figs 3436.
1976: Harpoceras (Harpoceras) subexaratum BONARELLI GABILLY, p. 108.
1979: Harpoceras exaratum (YOUNG & BIRD) subexaratum (BONARELLI) MARIOTTI et al., pl. 1, fig. 10.
? 2002a: Harpoceras subexaratum (BONARELLI) MACCHIONI, p. 130, fig. 77.
v 2007: Harpoceras subexaratum BONARELLI GCZY & SZENTE, pl. 1, fig. 6.
2010: Harpoceras cf. subexaratum BONARELLI VENTURI et al., p. 277.

Material from the H. serpentinum Zone 5 internal PINNA 1969, pl. 1, fig. 18, and VENTURI & FERRI 2001, p.
moulds of different state of preservation. 181). H. subexaratum is typical of the Mediterranean localities,
Description Moderately involute, compressed form and closely allied to H. mediterraneum, of which is thought to
with suboval whorl-section. The umbilicus is undercut, the be a descendant in the upper H. pseudoserpentinum horizon. It
margin is rounded. The flanks are slightly convex with differs from the latter by being slightly more involute, and
rounded shoulder; the venter is slightly broad, low and possessing finer, less developed ribs (see VENTURI et al.
carinate with a high keel. No whole body chamber is pre- 2010, p. 277, 303). A few Gerecse specimens with some-
served. The ribs are fine, dense and moderately falcoid. The what more evolute coiling can be regarded as transitional
suture-line consists of wide and short E, wide and long L, forms between the two taxa.
and short U lobes. The wide ES is asymmetrically divided. Both the H. (Harpoceras) cf. mulgravium and the Harpoceras
Remarks The specimens agree well with the type sp. figured by PINNA (1968, pl. 4, fig. 3, pl. 5, fig. 9) seem to be
(MENEGHINI 18671881, pl. 4, figs 1ac, refigured by very close in size and morphology to the Gerecse specimen

Explanation to Plate XII

1 Hildaites levisoni (SIMPSON) (244.2011), Bnya-hegy A, Bed 46, O. douvillei horizon, D: 118, H: 30 (25.4%), W: 22 (73.3%), U:
60 (50.8%).
23 Cingolites picenus SASSAROLI et VENTURI (265.2011), Crocodile Section, Bed 112, H. pseudoserpentinum horizon, D: 100, H: 28
(28%), W: 24 (85.7%), U: ?50.
4 Hildaites aff. serpentiniformis BUCKMAN (252.2011), Kis-Gerecse, Bed 106, H. pseudoserpentinum horizon, D: 134, H: 43
(32%), W: 22 (51%), U: 62 (46.2%).
5 Hildaites serpentiniformis BUCKMAN (245.2011), Kis-Gerecse, Bed 103, H. pseudoserpentinum horizon, D: 112, H: 33 (29.4%), W:
18 (54.5%), U: 56 (50%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 31
Plate XII

Fragmenta Palaeontologica Hungarica 29, 2011

32 KOVCS, Z.
figured on Plate X: 12. H. mulgravium was recognized as a 33 and 34), and the styles of ribbing are very similar.
synonym of H. falciferum by HOWARTH (1992), and this Consequently, the specimens in question might be regarded
arrangement was confirmed by BCAUD (2006). PINNAs as large examples of H. subexaratum. Osperleioceras (Pseudopoly-
specimens differ from the type of A. mulgravius (refigured by plectus) bicarinatum (ZIETEN) is a very similar form, but differs
BUCKMAN 1909, pls 4AB, and HOWARTH 1992, pl. 20, fig. in more involute coiling with ventral sulci and wider ribs.
5) by being more involute, lacking lateral groove and Distribution Italy, Greece, Montenegro, North
possessing finer ribs; on the other hand, they strongly Africa, South America. Hungary: Bakony Mts; Gerecse
resemble H. subexaratum in morphology. The suture-lines of Mts: Uppermost H. pseudoserpentinum horizon H. bifrons
both species are identical (see PINNA 1968, pl. 1 n.t., figs 13, Zone: Kis-Gerecse, Nagy-Pisznice and Bnya-hegy B
16, 19 and fig. 15), the whorl-sections (l.c., pl. 2 n.t., figs 29, Sections; O. douvillei horizon: Crocodile Section.

Harpoceras (Harpoceras) subplanatum (OPPEL, 1856)

(Plate VIII: 34, Plate XI: 3)
1846: Ammonites complanatus BRUGUIRE DORBIGNY, p. 353, pl. 114, figs 12, 4 (non fig. 3).
1856: Ammonites subplanatus OPPEL, p. 244.
1874: Ammonites subplanatus OPPEL DUMORTIER, p. 51, pl. 10, pl. 11, figs 12, 8.
1966: Polyplectus subplanatus (OPPEL) NUTSUBIDZE, p. 94, pl. 20, fig. 4, pl. 21, fig. 1.
v 1967b: Polyplectus subplanatus (OPPEL) GCZY, p. 123, text-fig. 6, pl. 2, fig. 4.
1986: Polyplectus subplanatus (OPPEL) GAKOVI, p. 115, pl. 19, fig. 2 (cum syn.).
1987: Polyplectus cf. subplanatus (OPPEL) HALL, p. 1698, pl. 3, figs AC.
1987: Harpoceras subplanatum (OPPEL) HILLEBRANDT, pl. 7, figs 35.
1992: Harpoceras subplanatum (OPPEL) HOWARTH, p. 136, text-figs 18H, 29, 35, 36, pl. 22, figs 47, pl. 23, figs 13 (cum syn.).
1994: Harpoceras subplanatum (OPPEL) GABILLY & MOUTERDE, p. 101, pl. 32, figs 1, 2.
2006: Harpoceras (Harpoceras) subplanatum (OPPEL) BCAUD, p. 65, pl. 25, fig. 1, pl. 31, fig. 2 (cum syn.).
? 2006: Harpoceras subplanatum (OPPEL) TOPCHISHVILI et al., pl. 13, figs 46.
? 2007: Harpoceras subplanatum (OPPEL) STEVENS, p. 379, text-figs 35.
v 2007: Harpoceras cf. subplanatum (OPPEL) GALCZ et al., p. 345, fig. 3/7.
2007: Harpoceras subplanatum (OPPEL) RULLEAU, p. 68, pl. 17, fig. 4, pl. 19, figs 12.

Material 2 moderately preserved internal moulds. Middle Toarcian by JAKOBS et al. (1994), and JAKOBS (1997),
Description Large, discoidal, involute form. The and from the H. falciferum Zone by DEAN et al. (1961),
umbilicus is moderately shallow, the umbilical wall is HALL (1987), and POULTON et al. (2005). The early occur-
undercut, the margin is rounded. The flanks are slightly rence of H. subplanatum in the Gerecse successions confirms
convex; the venter is narrow and carinate with a high the extended stratigraphic range. On the contrary, the very
keel. The whorl-section is lanceolate with maximum thick- early appearance of Polyplectus subplanatus (OPPEL) in the
ness at the mid-height. The body chambers are missing. upper P. hawskerense Subzone or D. tenuicostatum Zone
The ornamentation consists of simple, fine, dense, falcate (WIEDENMAYER 1980, p. 97, pl. 16, figs 1718) seems to be
ribs, which rise from the umbilicus, and fade at the keel. dubious. Eleganticeras elegans (SOWERBY) is a similar form in
The intricate suture-line corresponds to that figured by size and morphology, as well as it persisted from the upper
HOWARTH (1992, text-fig. 18/H). E. elegantulum to the upper H. falciferum Subzones; it
Remarks Both specimens agree well with the type markedly differs, however, in oblique and flat umbilical wall.
(DORBIGNY 1846, pl. 114, figs 12, 4, refigured by The classification of the species as Polyplectus has been rejected,
HOWARTH 1992, text-fig. 35CD). The species probably as the latter is an oxycone form with an acutely angled venter,
derived from H. falciferum in the upper H. pseudoserpentinum and the suture-line contains three or more auxiliary saddles.
horizon. The morphology of both taxa is similar, but H. Distribution World-wide. Hungary: Bakony Mts,
subplanatum is characterised by a more involute coiling with (?)Mecsek Mts; Gerecse Mts: Upper H. pseudoserpentinum
narrower whorls and finer ribs. Its range is typical of the H. horizon: Kis-Gerecse; O. douvillei horizon: Nagy-Pisznice
bifrons Zone, but it was also recorded from the Lower to Sections.

Explanation to Plate XIII

1 Hildaites murleyi (MOXON) (248.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 84, H: 24 (28.5%), W: 16 (66.6%), U: 40
(47.6%).
23 Hildaites subserpentinus BUCKMAN (247.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 70, H: 24 (34.2%), W: 12 (50%),
U: 26 (37%).
4 Hildaites levisoni (SIMPSON) (253.2011), Kis-Gerecse, Bed 101, O. douvillei horizon, D: 210, H: 58 (27.6%), W: 33 (56.8%), U:
108 (51.4%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 33
Plate XIII

Fragmenta Palaeontologica Hungarica 29, 2011

34 KOVCS, Z.
Genus Ovaticeras BUCKMAN, 1918
Ovaticeras cf. ovatum (YOUNG & BIRD, 1822)
(Plate IX: 3)
1822: Ammonites ovatus YOUNG & BIRD, p. 251, pl. 13, fig. 4.
1884: Harpoceras ovatum (YOUNG & BIRD) WRIGHT, p. 446, pl. 63, figs 47.
1918: Ovaticeras pseudovatum (YOUNG & BIRD) BUCKMAN, pls 111B, 111C.
1991: Ovaticeras sp. cf. O. ovatum (YOUNG & BIRD) POULTON, p. 20, pl. 13, figs 7, 11.
1992: Ovaticeras ovatum (YOUNG & BIRD) HOWARTH, p. 141, pl. 23, fig. 4, pl. 24, figs 14, pl. 25, figs 12 (cum syn.).
? 1992: Ovaticeras ovatum YOUNG & BIRD JIMNEZ & RIVAS, p. 49, pl. 1, fig. 9.
2005: Ovaticeras cf. ovatum (YOUNG & BIRD) POULTON et al., pl. 5, fig. 13.

Material 1 poorly preserved internal mould. wide and asymmetrical divided, the LS1 is long.
Description Large, moderately involute form Remarks The specimen resembles the type (YOUNG
without umbilical margin. The umbilicus is moderately & BIRD 1822, pl. 13, fig. 4, refigured by BUCKMAN 1918,
deep, the umbilical wall is rounded. The flanks are convex; pl. 111A, and HOWARTH 1992, pl. 24, fig. 4), and the
the venter is narrow and rounded, bearing a strong keel specimens figured in the literature. The absence of well-
on the inner whorls. The whorl-section is compressed developed keel on the last whorl of the figured specimen
elliptical. The body chamber is missing. The ornamenta- is due to the poor state of preservation.
tion is characterised by weakly developed, dense and Distribution England, France, Spain, North Africa,
falcoid ribs. The suture-line consists of wide E, long, North America. Gerecse Mts: O. douvillei horizon: Bnya-
narrow L, 2 accessory and 1 small U lobes. The ES is hegy B Section.

Subfamily Hildoceratinae HYATT, 1867

Genus Hildaites BUCKMAN, 1921
Hildaites wrighti (SPATH, 1913)
(Plate XV: 5, Plate XVI: 34)
1884: Harpoceras normaniaum (DORBIGNY) WRIGHT, p. 470, pl. 83, figs 12.
1909: Hildoceras serpentinum REINECKE RENZ, p. 222, pl. 4, fig. 2.
1913: Protogrammoceras wrighti SPATH, p. 553.
1963: Hildaites serpentinus (REINECKE) ZANZUCCHI, p. 126, pl. 18, fig. 8 only.
1982: Hildaites (?) (cf. exilis VENTURI) VENTURI, p. 72, fig. 104.
1987: Hildaites striatus GUEX KAZAKOVA, p. 99, pl. 2, fig. 5.
2002b: Hildaites wrighti (SPATH) MACCHIONI, fig. 4/6, 9.
2006: Hildaites wrighti (SPATH) BCAUD, p. 76, pl. 32, fig. 1 (cum syn.).
2009: Hildaites cf. striatus GUEX EL HAMMICHI et al., pl. 1, fig. 4.

Material 3 moderately preserved internal moulds. differs in finer and denser ribs (see below). Based on the
Description Evolute, compressed form with high morphology (style of coiling, ventral part, sculpture), five
suboval whorl-section. The umbilicus is wide and shallow, specimens are regarded here as H. wrighti from the
the margin is low and rounded, the flanks are slightly literature: H. serpentinus (RENZ 1909, p. 222, pl. 4, fig. 2),
convex, the shoulder is rounded. The venter is narrow H. serpentinus (ZANZUCCHI 1963, pl. 18, fig. 8 only), H.
and tricarinate-bisulcate with a low keel and shallow sulci. striatus (KAZAKOVA 1987, pl. 2, fig. 5), H. cf. striatus (EL
The ornamentation consists of moderately developed, HAMMICHI et al. 2009, pl. 1, fig. 4), and H. (?) (cf. exilis
simple, rectiradiate and slightly sigmoid ribs, which rise VENTURI) (VENTURI 1982, fig. 104). H. wrighti is a rare
from the umbilicus, and become projected on the ventro- but widespread species in both the Submediterranean and
lateral part. The suture-line is simple hildoceratid. the Mediterranean Provinces.
Remarks The specimens agree well with the type Distribution England, France, Spain, Italy, Greece,
(WRIGHT 1884, pl. 83, figs 12, refigured by HOWARTH North Africa, Caucasus. Gerecse Mts: H. striatus horizon:
1992, pl. 30, fig. 7). H. striatus GUEX is a similar form, but Kis-Gerecse, Nagy-Pisznice and Bnya-hegy B Sections.

Explanation to Plate XIV

12 Hildaites murleyi (MOXON) (254.2011), Nagy-Pisznice, Bed 134, O. douvillei horizon, D: 128, H: 37 (29%), W: 24 (64.8%), U: 58
(45.3%).
3 Hildaites forte (BUCKMAN) (251.2011), Kis-Gerecse, Bed 112, H. striatus horizon, D: appr. 76, H: appr. 20, W: ?, U: 42 (55.2%).
4 Hildaites crassus (GUEX) (246.2011), Kis-Gerecse, Bed 104, H. pseudoserpentinum horizon, D: 64, H: appr. 23 (36%), W: 20 (87%),
U: appr. 45 (70%).
56 Hildaites aff. pseudolevisoni VENTURI (255.2011), Kis-Gerecse, Bed 103, H. pseudoserpentinum horizon, L: 87, H: 26, W: 17
(65.3%).
78 Hildaites sp. (250.2011), Kis-Gerecse, Bed 106, H. pseudoserpentinum horizon, D: 106, H: 35 (33%), W: 18 (51.4%), U: 47 (44.3%).
Fragmenta Palaeontologica Hungarica 29, 2011
 Lower Toarcian Ammonitida 35
Plate XIV

Fragmenta Palaeontologica Hungarica 29, 2011

36 KOVCS, Z.
Hildaites striatus GUEX, 1973
(Plate XI: 2)
? 18671881: Ammonites serpentinus (REINECKE) MENEGHINI, p. 13, pl. 3, fig. 1.
1963: Hildaites serpentinus (REINECKE) ZANZUCCHI, p. 126, pl. 18, figs 1, 3, 67 only.
? 1969: Hildaites serpentinum (REINECKE) PINNA, p. 11, pl. 1, fig. 16.
1973: Hildaites striatus sp. n. GUEX, p. 504, pl. 2, fig. 5, pl. 3, fig. 10, pl. 4, fig. 1, pl. 7, fig. 3, pl. 9, fig. 2, pl. 10, fig. 2, pl. 14, fig. 11, 15, pl. 15, fig. 6.
1980: Hildaites striatus GUEX WIEDENMAYER, p. 100, pl. 32, figs 15.
1992: Hildaites striatus GUEX JIMENEZ & RIVAS, p. 60, pl. 4, figs 1115, pl. 5, figs 12.
2010: Hildaites striatus GUEX BILOTTA et al., fig. 17/AB.
2011: Hildaites striatus GUEX GALCZ et al., p. 326, pl. 3, fig. 1.

Material 4 moderately preserved internal moulds Remarks The specimens agree well with the type
and 6 fragmentary specimens. (GUEX 1973, pl. 7, fig. 3, pl. 14, fig. 11), and with the
Description Evolute form with narrow oval whorl- specimens figured in the literature. The species is
section. The umbilicus is wide and shallow, the margin is considered as the Mediterranean counterpart of H. wrighti
rounded. The flanks are slightly convex; the carinate (SPATH), from which it differs by bearing denser and
venter is narrow and low, and bisulcate on the inner finer ribs.
whorls. The ornamentation consists of dense, fine and Distribution Italy, southern Spain, Portugal, North
sinuous ribs. The suture-line is simple hildoceratid with Africa, (?)Greece, (?)Caucasus. Gerecse Mts: H. striatus
short and narrow E, wide L, and small, straight U lobes. horizon: all studied Gerecse sections.

Hildaites murleyi (MOXON, 1841)

(Plate XIII: 1, Plate XIV: 12, Plate XV: 12)
1841: Ammonites Murleyi MOXON, pl. 24, fig. 6.
1992: Hildaites murleyi (MOXON) HOWARTH, p. 168, pl. 30, figs 910, pl. 31, figs 18, pl. 32, fig. 4 (cum syn.).
2002b: Hildaites murleyi (MOXON) MACCHIONI, fig. 4/3, 10.
2006: Hildaites murleyi (MOXON) BCAUD, p. 77, pl. 32, fig. 3, pl. 33, fig. 1, pl. 34, fig. 1 (cum syn.).
2008: Hildaites murleyi (MOXON) METODIEV, fig. 4/k, l.

Material 27 inner moulds in various preservation. fig. 10), and to those figured in the literature. The species
Description Large, evolute form with sub-rectan- shows remarkable variability in morphology (HOWARTH
gular to broad subtrapezoidal whorl-section. The umbili- 1992). According to HOWARTH (l.c.), and BCAUD (2006),
cus is wide and shallow, the margin is rounded, the flanks Hildaites borealis (SEEBACH), H. propeserpentinus (BUCKMAN)
are flat, the shoulder is pronounced. The ventral part is and Murleyiceras aptum BUCKMAN are synonyms of H.
carinate and bisulcate on the inner whorls, while flat with murleyi, consequently the abundance of the species in the
a keel on the body chamber. The ornamentation consists Mediterranean Province is also well-documented. The style of
of mostly simple, rursiradiate, sinuous and coarse ribs. ribbing of the specimen figured on Plate XIII: 1 resembles
The ribs rise from the margin, curve back on the dorsal that of H. eremitensis VENTURI (1973, figs 5, 13/e, h). H. murleyi
half, then forward on the ventral half of the whorl, and is closely allied to H. undicosta (MERLA) in morphology, but
reach the shoulder. Some ribs bifurcate irregularly at the differs by stronger and more rursiradiate ribs.
margin. The suture-line is simple with short E, wide and Distribution Europe, North Africa, Greenland,
slightly longer L, and small, straight U lobes. Caucasus, northern Russia, South and North America.
Remarks The specimens are close to the type Gerecse Mts: H. striatus horizon: Crocodile Section; H.
(MOXON 1841, pl. 24, fig. 6, refigured by BUCKMAN striatus H. pseudoserpentinum horizons: Kis-Gerecse, Bnya-
1921, pl. 216, figs 13, and by HOWARTH 1992, pl. 30, hegy B and Nagy-Pisznice Sections.

Hildaites forte (BUCKMAN, 1921)

(Plate XIV: 3)
1921: Murleyiceras forte BUCKMAN, pl. 245.
1976: Hildoceras (Hildaites) levisoni (SIMPSON) SCHLEGELMILCH, p. 85, pl. 44, fig. 3.
1992: Hildaites forte (BUCKMAN) HOWARTH, p. 171, pl. 32, figs 13, text-figs 4041 (cum syn.).

Explanation to Plate XV
12 Hildaites murleyi (MOXON) (262.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 46, H: 14 (30.4%), W: ?, U: 26 (56.5%).
34 Orthildaites douvillei (HAUG) morphotype raricostata (MITZOPOULOS) (259.2011), Bnya-hegy A, Bed 46, O. douvillei
horizon, D: 86, H: 22 (25.5%), W: 20 (91%), U: 44 (51%).
5 Hildaites wrighti (SPATH) (249.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 105, H: 30 (28.5%), W: 16 (53.3%), U: 50 (47.6%).
67 Orthildaites douvillei (HAUG) (258.2011), Kis-Gerecse, Bed 102, O. douvillei horizon, L: 95, H: 27, W: 23 (85%), U: ?.
8 Hildaites serpentiniformis BUCKMAN (256.2011), Nagy-Pisznice, Bed 135, H. pseudoserpentinum horizon, D: 154, H: 45 (29.2%),
W: 22 (48.8%), U: 76 (49.3%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 37
Plate XV

Fragmenta Palaeontologica Hungarica 29, 2011

38 KOVCS, Z.
? 2004: Hildaites levisoni (SIMPSON) MORARD, pl. 19, fig. 7 only.
2006: Hildaites forte (BUCKMAN) BCAUD, p. 78, pl. 35, fig. 1 (cum syn.).
v? 2007: Hildaites forte (BUCKMAN) GALCZ et al., p. 345, fig. 3/9.
2008: Hildaites cf. forte (BUCKMAN) GALCZ et al., pl. 8, figs 12.
2008: Hildaites forte (BUCKMAN) METODIEV, fig. 4/m, n.

Material 1 fragmentary specimen. Remarks The specimen shows high similarity to

Description Evolute form with subquadrate the holotype (BUCKMAN 1921, pl. 245). According to
whorl-section. Both the margin and the low umbilical HOWARTH (1992), Murleyiceras gyrale BUCKMAN is a
wall are rounded. The flanks are flat and parallel. The synonym of H. forte. H. murleyi is a closely allied form, but
venter is wide and flat, and tricarinate-bisulcate. No differs in denser, less-developed ribbing, and in higher
whole body chamber is preserved. The ribs are simple, whorl-section.
rursiradiate, coarse, and are projected on the ventral Distribution Europe, North Africa, Iran. Hungary:
half of the flank. The intercosta is wider than the rib- (?)Bakonycsernye; Gerecse Mts: H. striatus horizon: Kis-
width. The suture-line is similar to that of H. murleyi. Gerecse Section.

Hildaites levisoni (SIMPSON, 1843)

(Plate XII: 1, Plate XIII: 4)
1843: Ammonites Levisoni SIMPSON, p. 54.
1910: Hildoceras levisoni SIMPSON BUCKMAN, pl. 12, figs 12.
1985: Hildoceras (Hildaites) levisoni levisoni (SIMPSON) RIEGRAF, p. 258, pl. 15, fig. 2, pl. 16, figs 1, 3.
1986: Hildaites cf. levisoni (SIMPSON) GAKOVI, p. 106, pl. 15, fig. 1ab.
2006: Hildaites levisoni (SIMPSON) BCAUD, p. 79, pl. 33, fig. 3, pl. 36, fig. 1 (cum syn.).
2007: Hildaites levisoni (SIMPSON) RULLEAU, p. 71, pl. 18, fig. 3.

Material 9 inner moulds of various preservation. morphology of the adult form, and the range of the species
Description Large, evolute form with sub-rectangular were widely discussed in the literature, and the determinations
whorl-section. Both the umbilical wall and the margin are showed uncertainty. Consequently, the taxon was included
rounded. The flanks are parallel with a moderately wide, within H. murleyi by HOWARTH (1992). The species was
concave, shallow lateral groove on the dorsal half. The ventral reinvestigated by BCAUD (2006: 7980), who provided a
part is wide, low and rounded, tricarinate-bisulcate on the detailed comparison with H. murleyi, and established again the
inner whorls, but the sulci disappear on the last whorl, and the validity of the taxon. The species differs from other Hildaites
venter becomes smooth and carinate. No whole body species in the marked lateral groove on the adult body
chamber is preserved. The ornamentation consists of broad, chamber. The Gerecse specimens are close to the large ones
sigmoid and slightly rursiradiate ribs with wider intercosta figured in the literature (e.g. RIEGRAF 1985, pl. 16, figs 13;
than the rib-width. The ribs are weakly developed on the GOY & MARTNEZ 1990, pl. 1, fig. 6; GOY et al. 1994, p. 113,
dorsal half, while projected on the ventral half of the flank. fig. 6; BCAUD 2006, pl. 33, fig. 3, pl. 36, fig. 1).
The suture-line is hildoceratid. Distribution Europe, North Africa, South America,
Remarks As the type is a small specimen (figured by eastern Russia. Hungary: Mecsek Mts; Gerecse Mts: H.
BUCKMAN 1910, pl. 12, refigured by SCHLEGELMILCH 1976, falciferum Subzone: Kis-Gerecse Section, O. douvillei horizon:
pl. 44, fig. 2, and HOWARTH 1992, pl. 31, fig. 2), both the Nagy-Pisznice and Crocodile Sections.

Hildaites subserpentinus BUCKMAN, 1921

(Plate XIII: 23)
1921: Hildaites subserpentinus BUCKMAN, pl. 217.
1963: Hildaites subserpentinus BUCKMAN KOTTEK, p. 74, text-fig. 27, pl. 5, figs 12.
1975: Hildaites subserpentinus BUCKMAN VENTURI, text-fig. 1/a, pl. 30, fig. 7.
1992: Hildaites subserpentinus BUCKMAN HOWARTH, p. 174, pl. 32, fig. 5, pl. 33, figs 14, pl. 34, fig. 1.
2001: Hildaites subserpentinus BUCKMAN VENTURI & FERRI, p. 196, figs fh.
2006: Hildaites subserpentinus BUCKMAN BCAUD, p. 80, pl. 35, fig. 2, pl. 37, fig. 1 (cum syn.).
2007: Hildaites subserpentinus BUCKMAN RULLEAU, p. 71, pl. 21, fig. 3.
2011: Hildaites subserpentinus BUCKMAN GALCZ et al., p. 326, pl. 1, fig. 3.

Explanation to Plate XVI

12 Orthildaites sp. A (260.2011), Nagy-Pisznice, Bed 133, O. douvillei horizon, D: 114, H: 30 (26.3%), W: 28 (93.3%), U: 59 (51.7%).
34 Hildaites wrighti (SPATH) (243.2011), Kis-Gerecse, Bed 113b, H. striatus horizon, D: 61, H: 20 (32.7%), W: 12 (60%), U: 27 (44.2%).
5 Hildaites cf. undicosta (MERLA) (264.2011), Bnya-hegy B, Bed 12, H. striatus horizon, D: 86, H: 27 (31.3%), W: 18 (66.6%), U: 40 (46.5%).
6 Orthildaites becaudi n. sp. (261.2011), paratype, Bnya-hegy A, Bed 46, O. douvillei horizon, D: 60, H: 16 (26.6%), W: 12 (75%), U: 29
(48.3%).
78 Orthildaites becaudi n. sp. (263.2011), holotype, Kis-Gerecse, Bed 102, O. douvillei horizon, D: 103, H: 24 (23.3%), W: 18 (75%), U: 53
(51.4%).

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 39
Plate XVI

Fragmenta Palaeontologica Hungarica 29, 2011

40 KOVCS, Z.
Material 20 internal moulds of different state of Remarks Based on the revision of genus Hildaites
preservation. accomplished by HOWARTH (1992), some Hildaites serpen-
Description Moderately evolute form with suboval tinus (REINECKE) records in the literature were reclassified
whorl-section. Both the umbilical wall and the margin are as H. subserpentinus BUCKMAN by BCAUD (2006: 80). The
rounded; the flanks are slightly convex. The venter is trica- Gerecse specimens agree well with the type (BUCKMAN
rinate-bisulcate on the inner whorls, while rounded and 1921, pl. 217, refigured by HOWARTH 1992, pl. 33, fig. 1).
carinate without ventral sulci on the external whorls. No The species differs from H. murleyi in suboval whorl-section.
whole body chamber is preserved. The ornamentation It is closely allied to the Mediterranean Hildaites exilis
consists of well-developed, somewhat dense, rursiradiate, VENTURI in morphology, but differs in stronger ribs.
sinuous ribbing. Mostly simple, strong ribs emerge from the Distribution Europe, North Africa, (?)North Ame-
umbilicus, become slightly projected on the outer half of the rica. Gerecse Mts: H. striatus horizon: Nagy-Pisznice, Croco-
whorl, and fade at the venter. The suture-line is simple with dile and Bnya-hegy B, H. striatus H. pseudoserpentinum
short E, moderately wide and long L, and small U lobes. horizons: Kis-Gerecse Sections.

Hildaites serpentiniformis BUCKMAN, 1921

(Plate XII: 5, Plate XV: 8)
1921: Hildaites serpentiniformis BUCKMAN, p. 55.
1923: Hildaites serpentiniformis BUCKMAN, pl. 267B.
V 1967a: Hildaites aff. serpentiniformis BUCKMAN GCZY, p. 128, text-fig. 126, pl. 30, fig. 2.
1987: Hildaites cf. serpentiniformis BUCKMAN HALL, p. 1700, pl. 4, figs AC.
1996: Hildaites aff. H. serpentiniformis urkutensis GCZY POPA & PATRULIUS, pl. 4, fig. 8.
2006: Hildaites serpentiniformis BUCKMAN BCAUD, p. 80, pl. 38, fig. 1 (cum syn.).

Material 8 internal moulds of different state of wide and longer L, and short U lobes.
preservation. Remarks The specimens are close to the type
Description Evolute, compressed form with (BUCKMAN 1923, pl. 267B, refigured by HOWARTH 1992,
shallow umbilicus. Both the umbilical wall and the margin pl. 33, fig. 4), and to the specimens figured in the litera-
are rounded; the flanks are flat. The venter is tricarinate- ture. They differ from H. serpentiniformis urkutensis GCZY
bisulcate on the inner whorls, while rounded and carinate by having slightly stronger ribs on the dorsal half of the
on the last whorl. The section is subtrapezoidal with external whorls. The species differs from H. murleyi, H.
maximum thickness at the margin. The preserved body undicosta, and H. subserpentinus by being less evolute with
chamber is half a whorl in length. The ornamentation subtrapezoidal section, by bearing weakly developed ribs on
consists of moderately developed, sigmoid ribs. Mostly the inner half of the whorl, and by having wider L lobe.
simple, weakly developed ribs rise from the umbilicus, Distribution Europe, North Africa, Caucasus, South
become projected on the outer half of the whorl, and and North America. Hungary: Bakony Mts; Gerecse Mts:
fade at the venter. Some rib-pairs emerge from umbilical uppermost H. striatus horizon: Bnya-hegy B, H. falciferum
nodes irregularly. The suture-line is simple with short E, Subzone: Kis-Gerecse and Nagy-Pisznice Sections.

Hildaites aff. serpentiniformis BUCKMAN, 1921

(Plate XII: 4)
Material 1 fragmentary specimen. projected on the ventrolateral part. The suture-line consists of
Description Large, evolute, compressed form with slightly asymmetric ES, long and wide L, asymmetric LS and
high, rounded subrectangular whorl-section. Both the high short U lobes.
umbilical wall and the margin are rounded; the flanks are Remarks Hildaites serpentiniformis BUCKMAN seems to
almost flat and parallel. The shoulder is rounded; the venter is be the most similar form in morphology, but differs from the
narrow, low and tricarinate-bisulcate. The first quarter of the specimen figured here in whorl-section and style of ribbing.
body chamber is preserved. The ribs are simple, rursiradiate Distribution H. pseudoserpentinum horizon of the
and falcoid, less developed on the dorsolateral, while Kis-Gerecse Section (Bed 106).

Hildaites crassus (GUEX, 1973)

(Plate XIV: 4)
1973: Mercaticeras crassum sp. n. GUEX, p. 506, pl. 6, fig. 1, pl. 14, fig. 23.
2001: Hildaites crassus (GUEX) VENTURI & FERRI, p. 196, figs ac.
v 2008: Hildaites crassus (GUEX) GALCZ et al., pl. 7, figs 56.
2010: Hildaites crassus (GUEX) VENTURI et al., p. 313.

Material 1 fragmentary specimen. are slightly convex, the venter is low and broad, tricarinate-
Description Evolute form with subquadrate whorl- bisulcate. The ornamentation is characterised by simple,
section. The umbilical wall is high and rounded, the flanks rursiradiate, wide and rounded, slightly sinuous ribs rising
Fragmenta Palaeontologica Hungarica 29, 2011
 Lower Toarcian Ammonitida 41
from the umbilicus. The suture-line is hildoceratid. line, however, ribs of H. crassus are not clavate, and are
Remarks This rare species is typical of the Tethyan coarser with a hildaitic rursiradiate-falcate appearance.
region. The specimen figured here shows a good agreement (SASSAROLI & VENTURI 2010).
with the holotype (GUEX 1973, pl. 6, fig. 1). The species is Distribution Italy, North Africa. Hungary: Bakony
closely allied to genus Cingolites SASSAROLI et VENTURI by Mts; Gerecse Mts: H. pseudoserpentinum horizon: Kis-
having a wide tricarinate-bisulcate venter and a simple suture Gerecse Section.

Hildaites cf. undicosta (MERLA, 1933)

(Plate XVI: 5)
1933: Hildoceras undicosta n. sp. MERLA, p. 50, pl. 7, fig. 2.
1947: Hildoceras undicosta MERLA LIPPI BONCAMBI, p. 138, pl. 2, figs 1314.
1963: Hildaites propeserpentinus (BUCKMAN) KOTTEK, p. 70, text-figs 2526, 58, pl. 4, figs 57, pl. 13, figs 45.
1973: Hildaites cf. undicosta (MERLA) VENTURI, p. 593, text-figs 8, 14/c.
non 1975: Hildaites undicosta (MERLA) VENTURI, pl. 30, fig. 8 (= H. pseudolevisoni VENTURI).
1980: Hildaites undicosta (MERLA) WIEDENMAYER, p. 101, pl. 32, figs 610.
2001: Hildaites undicosta (MERLA) VENTURI & FERRI, p. 192.
2003: Hildaites gr. undicosta VENTURI & ROSSI, p. 71.
2010: Hildaites undicosta (MERLA) VENTURI et al., p. 313.

Material 4 poorly preserved internal moulds. accepted in the Italian literature (although, without any
Description Evolute form with subrectangular detailed comparison with H. murleyi). Its range was also
whorl-section. Both the umbilical wall and the margin are recorded from the Btic Range (GONZLEZDONOSO et
rounded, the flanks are flat and parallel, the venter is al. 1971) and from the Bakony Mts (GCZY 1975). The
tricarinate-bisulcate. The ornamentation is characterised by species is considered as a Mediterranean counterpart of H.
slightly rursiradiate, simple, somewhat dense and sigmoid murleyi in this paper. The Gerecse specimens with subquad-
ribs. The suture-line is hildoceratid. rate section seem to be close to the type (MERLA 1933, pl.
Remarks The species is one of the characteristic 7, fig. 2), but differ from the more compressed ones pre-
Mediterranean Hildaites, abundant in Italian localities. It is sented by VENTURI et al. (2010: 313). Due to the poor
closely allied with H. murleyi in morphology, but differs by preservation, the exact determination cannot be arranged.
lower whorls, and slightly finer and regular ribs with less Distribution Italy, Greece, southern Spain.
rursiradite direction. It was recognised as one of the syno- Hungary: Bakony Mts; Gerecse Mts: H. striatus horizon:
nyms of H. murleyi by HOWARTH (1992), and BCAUD Bnya-hegy B, O. douvillei horizon: Nagy-Pisznice Sec-
(2006); the validity of H. undicosta, however, is widely tions.

Hildaites aff. pseudolevisoni VENTURI, 1981

(Plate XIV: 56)
1981: Hildaites pseudolevisoni n. sp. VENTURI, p. 594, text-fig. 6, fig. 7/6ab (cum syn.).
1982: Hildaites pseudolevisoni VENTURI VENTURI, p. 71, fig. 102, p. 72, fig. 103.
1994: Hildaites pseudolevisoni VENTURI FARAONI et al., pl. 13, figs 2324.
1998: Hildaites pseudolevisoni VENTURI PARISI et al., text-figs 7/16, 8/1, pl. 3, fig. 14.
2001: Hildaites pseudolevisoni VENTURI VENTURI & FERRI, p. 195, fig. e.

Material 3 fragmentary internal moulds. and fade at the shoulder. The suture-line consists of short
Description Moderately evolute form with E, short and wide L, and small U lobes with broad LS
subtrapezoid whorl-section. The umbilical wall is steep and divided by a short and wide accessory lobe.
flat, the margin is rounded. The flanks are slightly convex, Remarks The specimen figured here differs from
the shoulder is rounded. The venter is narrow and low, and H. levisoni in finer ribs and absence of lateral groove; and
tricarinate-bisulcate. No whole body chamber is preserved. from H. subserpentinus in less sigmoid, widely spaced ribs.
The ornamentation consists of well-developed, simple and H. pseudolevisoni seems to be the closest form; however, it
sinuous ribs with slightly wider intercosta than the rib- differs in narrower intercosta.
width. The ribs emerge from the margin, become stronger Distribution Italy. Gerecse Mts: H. falciferum
at the mid-height, projected on the outer half of the flank, Subzone: Kis-Gerecse and Nagy-Pisznice Sections.

Hildaites sp.
(Plate XIV: 78)
Material A fragmentary wholly septate inner mould. rounded. The venter is moderately high, rounded, and
Description Evolute form with shallow umbilicus. carinate with a narrow and low keel without sulci. The
The umbilical wall is high, oblique and flat, the margin is whorl-section is suboval with maximum width under the
rounded. The flanks are slightly convex, the shoulder is mid-flank. The ornamentation is characterised by simple,

Fragmenta Palaeontologica Hungarica 29, 2011

42 KOVCS, Z.
radial, sigmoid ribs. They emerge from the margin, become Remarks The suboval whorl-section is typical of H.
projected on the ventrolateral part, and disappear on the subserpentinus, but the latter differs in denser ribs. The speci-
venter without reaching the keel. The intercosta is concave, men probably represents a new Mediterranean Hildaites
and wider than the rib-width. The hildoceratid suture-line form, but the arrangement needs further comprehensive
consists of narrow E and broad L with equal length, wide comparisons.
ES, and wide, asymmetrical divided LS. The U lobes Distribution H. pseudoserpentinum horizon of the
cannot be traced. Kis-Gerecse Section (Bed 106).

Genus Orthildaites BUCKMAN, 1923

Orthildaites douvillei (HAUG, 1884)
(Plate XV: 34, 67)
1884: Harpoceras Douvillei HAUG, p. 353, pl. 15, figs 1ac.
v 1967b: Hildoceras sublevisoni raricostatum MITZOPOULOS GCZY, p. 128, pl. 3, fig. 3.
1971: Orthildaites cf. orthus BUCKMAN GCZY, p. 49, fig. 14.
1976: Orthildaites douvillei (HAUG) GABILLY, p. 120, pl. 19, figs 113, pl. 20, figs 15 (cum syn.).
1976: Orthildaites douvillei (HAUG) morphotype raricostata (MITZOPOULOS) GABILLY, p. 125, pl. 20, figs 13.
? 1979: Orthildaites douvillei (HAUG) MARIOTTI et al., pl. 1, fig. 12.
2001: Orthildaites douvillei (HAUG) VENTURI & FERRI, p. 193, 196, figs jo only.
2006: Orthildaites orthus (BUCK.) TOPCHISHVILI et al., 2006, pl. 16, fig. 4.
2006: Orthildaites douvillei (HAUG) BCAUD, p. 83, pl. 40, fig. 2, pl. 42, fig. 1 (cum syn.).
2010: Orthildaites douvillei (HAUG) VENTURI et al., p. 314.

Material 17 internal moulds of different state of BUCKMAN, 1923 is a junior synonym of O. douvillei. The
preservation. latter shows slight variabilities in width of the whorls, and
Description Evolute, robust form with rounded shape and width of the ribs. Most Gerecse specimens with
subquadrate whorl-section. The umbilicus is wide and subquadrate section and broad, widely spaced ribs agree
shallow, the umbilical wall is steep and flat, the margin is perfectly with the type (HAUG 1884, pl. 15, figs 1ac,
rounded. The flanks are slightly convex, the shoulder is refigured by GABILLY 1976, pl. 19, figs 14) in morpho-
rounded. The venter is broad, low and carinate with a logy (Plate XV: 67). The specimen figured on Plate XV:
strong keel, and bears two wide and shallow grooves. No 34 represents O. douvillei morphotype raricostata (MITZO-
whole body chamber is preserved. The ornamentation POULOS). Hildoceras sublevisoni and H. caterinii MERLA are
consists of well-developed, simple, gently curved and similar forms, but differ in specific rursiradiate ribs on the
rectiradiate ribs with wider intercosta than the rib-width. innermost whorls, and in shorter, narrower and straight
The ribs rise from the margin, are straight on the inner ribs on the last whorl.
whorls, but widely spaced and slightly curved on the Distribution Europe, North Africa, Caucasus,
external whorls. The suture-line is simple with short E, North America. Hungary: Bakony Mts; Gerecse Mts: O.
broad and short L, and small, straight U lobes. douvillei horizon: Kis-Gerecse, Nagy-Pisznice and Croco-
Remarks According to HOWARTH (1992), and dile, O. douvillei horizon H. sublevisoni Subzone: Bnya-
BCAUD (2006), the type species of the genus, O. orthus hegy B Sections.

Orthildaites sp. A
(Plate XVI: 12)
Material 1 well-preserved internal mould and a uncommon in the Subboreal Submediterranean regions,
fragmentary specimen. however, similarly to Hildaites, more abundant and
Description Evolute form with subrectangular diversified in the Mediterranean Province (GUEX 1973;
whorl-section. The umbilical wall is oblique and flat, BILOTTA et al. 2010; SASSAROLI & VENTURI 2010).
the margin is rounded, the flank is slightly convex. The The specimen figured on Plate XVI: 12 markedly
venter is broad, low, tabulate and carinate. The pre- differs from O. douvillei in higher whorls with subrect-
served body chamber of the figured specimen is half a angular section, and in denser, narrower and somewhat
whorl long. The ornamentation is characterised by strong, sigmoid ribs. It resembles well with the O. douvillei
rectiradiate and straight ribs on the inner whorls, while specimens figured by MACCHIONI & VENTURI (1996,
slightly sigmoid ribs on the last whorl. The ribs rise pl. 2, figs 12) in morphology, and represents a new
from the margin, and fade away at the shoulder. The Orthildaites species (pers. inf. by Prof. F. V ENTURI and
simple suture-line agrees with that of O. douvillei. S. SASSAROLI).
Remarks According to HOWARTH (1992), Distribution Italy. Gerecse Mts: O. douvillei horizon:
Orthildaites is a monospecific genus. In reality, it is Kis-Gerecse and Nagy-Pisznice Sections.

Fragmenta Palaeontologica Hungarica 29, 2011

 Lower Toarcian Ammonitida 43
Orthildaites becaudi n. sp.
(Plate XVI: 68)
Holotype 263.2011, Natural History Museum of the Faculty of Sciences of Etvs Lornd University.
Paratype 261.2011, Natural History Museum of the Faculty of Sciences of Etvs Lornd University.
Derivation of name In honour of Marc Bcaud, the late French paleontologist.
Type horizon and locality O. douvillei horizon, Bed 102, Kis-Gerecse Section, Gerecse Mts.
Diagnosis Evolute form, wide and shallow umbilicus, low umbilical wall, flat and parallel lateral walls, tricarinate-
bisulcate venter. High subrectangular whorl-section, rectiradiate and slightly sigmoid ribs, hildoceratid suture-line.
Material 1 moderately preserved internal mould and a fragmentary specimen.
Measurements D H H/D W W/H U U/D
holotype 103 24 23.3% 18 75% 53 51.4%
paratype 60 16 26.6% 12 75% 29 48.3%
Description Evolute, moderately compressed form and denser ribs clearly distinguish the new species from
with high subrectangular whorl-section. The umbilical wall both O. douvillei and O. intermedius GUEX. It differs from
is low and steep, the flanks are parallel, the venter is tri- Orthildaites sp. A (Plate XVI: 12) in appearance of the
carinate-bisulcate. No whole body chamber is preserved. ventral part, and in style of ribbing. O. douvillei raricostata
The ornamentation is characterised by simple, rectiradiate (Plate XV: 34) is similar in ornamentation, but it is a
and slightly sigmoid ribs. The ribs emerge from the umbili- more robust form with lower and broader whorls (see
cus, and become projected on the ventrolateral part. The MITZOPOULOS 1930 and GABILLY 1976). Hildoceras
suture-line is hildoceratid with short E, broad and short L, caterinii differs by bearing rursiradiate ribs on the inner
and small, straight U lobes. whorls.
Remarks The specimens figured here are Distribution O. douvillei horizon: Kis-Gerecse and
arranged within genus Orthildaites on the basis of radial Bnya-hegy A Sections. The paratype came from Bed 46
inner ribs. They differ from other Orthildaites species of the Bnya-hegy A Section, associated with Orthildaites
described in the literature by well-defined morphological douvillei, Hildaites levisoni, Harpoceras mediterraneum, H.
features. The narrower whorls and somewhat sharper subexaratum, and Nodicoeloceras tuberculatum.

Genus Cingolites SASSAROLI et VENTURI, 2010

Cingolites picenus SASSAROLI et VENTURI, 2010
(Plate XII: 23)
2010: Cingolites picenus n. sp. SASSAROLI & VENTURI, p. 106, text-fig. 6/af, il, n, pl. 1, figs 58, pl. 2, figs 15, 9.
2010: Cingolites gr. picenus SASSAROLI & VENTURI, p. 108, text-fig. 6/gh, m, pl. 2, figs 67.
2011: Cingolites picenus SASSAROLI et VENTURI GALCZ et al., p. 327, pl. 1, figs 1-2.

Material 1 poorly preserved internal mould. the last one is widespread in the Hildoceras bifrons
Description Evolute form with subquadrate whorl- Zone of the Mediterranean area (recently figured from
section. The umbilical wall is vertical, the margin is the Gerecse Mts by GCZY & SZENTE 2007, pl. 4, fig.
rounded, the flank is slightly convex. The venter is broad, 3). According to the authors, Cingolites is distinguished
low and tricarinate-bisulcate with wide and deep sulci. from genus Orthildaites in sinuous and projected, clavate
The preserved body chamber is about half a whorl long. ribs, and coronate, strongly tricarinate-bisulcate ventral
The ornamentation consists of simple, rounded, clavate, part. It also differs from Mercaticeras in larger size,
and slightly rursiradiate ribs with wide, concave inter- coarser ornamentation, wider whorls, and in stratigraphic
costa. The ribs become gently sigmoid on the last whorl. distribution. The specimen figured here agrees with the
The hildoceratid suture-line is rather simple with short E, holotype of C. picenus (SASSAROLI & VENTURI 2010, pl. 1,
broad L, and small, straight U lobes. fig. 7) in morphology. The occurrence of the species in
Remarks Genus Cingolites was introduced by SASSA- the Gerecse assemblage confirms again the close rela-
ROLI & VENTURI (2010) with four species: C. clavatus n. tionship between the Italian and Hungarian Toarcian
sp., C. picenus n. sp., C. spiralis n. sp. and C. stefaninii ammonite faunas.
(MERLA). The first three species range in the upper Distribution Italy. Gerecse Mts: H. pseudoserpentinum
Undicosta/Falciferum Subzone in Italian localities; while horizon of the Crocodile Section (Bed 112).

Conclusions
The revision of the ammonite assemblages of the Crocodile Section provided a detailed picture of the Lower
classic Kis-Gerecse, Nagy-Pisznice, and Bnya-hegy AB Toarcian ammonite fauna of the Gerecse Mts. Based on
Sections, as well as the investigation of the newly excavated the general features of the assemblage, it can be placed
Fragmenta Palaeontologica Hungarica 29, 2011
44 KOVCS, Z.
between the Mediterranean ones known from Italian, taxa typical of NW European areas (Harpoceras serpentinum, H.
Southern Spanish, and Austrian localities on one hand, and strangewaysi, H. falciferum, H. pseudoserpentinum) made it possible
the Submediterranean ones recorded from France, Northern to establish the Submediterranean ammonite biostratigraphy.
Spain, and Bulgaria on the other. The composition of the The Dactylioceras tenuicostatum Zone can be demonstrated
Gerecse fauna shows a clear Tethyan character with domi- only from the lowermost part of the Kis-Gerecse Section,
nance of suborders Phylloceratina and Lytoceratina, with while strata of the Harpoceras serpentinum Zone are
occurrence of characteristic Mediterranean Ammonitina explored in most Toarcian Gerecse sections. For the latter
species (e.g. Mesodactylites mediterraneus, Nodicoeloceras tubercu- zone a slightly simplified stratigraphic scheme with two
latum, Harpoceras mediterraneum, H. subexaratum, Cingolites picenus), subzones (Harpoceras serpentinum, Harpoceras falciferum)
and with high diversity of genera Hildaites and Orthildaites. and three biohorizons (Hildaites striatus, Harpoceras pseudoserpen-
However, occurrences of important zonal index Ammonitina tinum, Orthildaites douvillei) are recognized here.

***

Acknowledgements I am grateful to Barnabs GCZY, Andrs GALCZ, Mikls KZMR and Istvn SZENTE (Etvs Lornd University,
Budapest), Istvn FZY (Hungarian Natural History Museum), Lszl KORDOS (Geological Institute of Hungary), Zoltn EVANICS (Budapest),
Soledad URETA (Madrid), Louis RULLEAU (Lyon), Massimiliano BILOTTA and Stefano SASSAROLI (Rosora) for their professional help. I express
my special acknowledgments to Prof. Federico VENTURI (Perugia) for his previous critical notes on my determinations of some Hildaites and
Orthildaites species.

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Authors address:
KOVCS Zoltn
Liszt Ferenc University
1076 Budapest, Liszt Ferenc tr 8.
Hungary
E-mail: [email protected]

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