AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE FROM

Transcription

Geologica Romana 40 (2007), 1-19
AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE FROM
PERI-MEDITERRANEAN TETHYAN SEDIMENTS (CALABRIA, SOUTHERN ITALY)
Nino Mariotti°, Massimo Santantonio° & Robert Weis°°
° Dipartimento di Scienze della Terra - Università “La Sapienza” - Piazzale Aldo Moro, 5 - 00185 Rome, Italy
[email protected]; [email protected]
°° Robert Weis - Musée National d’Histoire Naturelle de Luxembourg - Section Paléontologie
25, rue Münster - L-2160 Luxembourg - [email protected]
ABSTRACT - A belemnite fauna was collected in southern Italy from hemipelagic marls (Sant’Onofrio formation) of Aalenian-Early Bajocian age. This is part of the Jurassic-Lower Cretaceous Caloveto Group succession,
cropping out near the village of Caloveto (Calabria, Sila Mts., southern Italy). The cephalopod assemblage was
analyzed for systematics and biostratigraphy, and for highlighting its palaeobiogeographic significance.
Age assignments were constrained through cross-check with ammonite assemblages (Tmetoceras scissum,
Ancolioceras opalinoides, Erycites fallifax, Erycites intermedius, Docidoceras sp.) found in the belemnite-bearing levels.
The belemnite fauna is mainly composed of taxa ascribed to the genus Holcobelus (Early Aalenian-Middle
Bajocian). This genus is characterized by an apical groove extending from the apex almost reaching the alveolar
region. Although the species Holcobelus trauthi, Holcobelus subblainvillei, Holcobelus munieri, Holcobelus
tetramerus and Holcobelus tschegemensis are present, the majority of the specimens cannot be identified as
belonging to Holcobelus since their feature set also includes a very short apical region, a long and wide ventral
groove, and a post alveolar compression. These characters are found separately in belemnites belonging to different families (Megateuthididae and Pachybelemnopseidae). For this reason we maintain an open nomenclature
(Belemnitida incertae sedis) for these specimens, for which more systematic work is needed. Besides Holcobelus
and allied forms, the following taxa were also identified: Megateuthis sp., Brevibelus breviformis, Pachybelemnopsis baculiformis, Hibolithes wuerttembergicus and Hibolithes sp. This fauna has close affinities to faunas from Romania, Bulgaria, France, southern Germany, Luxembourg, England and Caucasus.
KEY WORDS: Belemnites, biostratigraphy, Calabria, Aalenian-Early Bajocian.
INTRODUCTION
PREVIOUS WORK
The purpose of this paper is to describe in detail a
peculiar belemnite fauna, composed of taxa ascribed to
Holcobelus Stolley, 1927, Megateuthis Bayle, 1878,
Brevibelus Doyle, 1992, Pachybelemnopsis Riegraf,
1980, Hibolithes Denis de Montfort, 1808 and
Belemnitida incertae sedis. The belemnites were collected from Aalenian-Early Bajocian red marls of the
Sant’Onofrio formation (Caloveto Group) outcropping
between the 8,2 and 8,4 km of the road from Caloveto to
Bocchiglierio villages (Calabria, southern Italy), in a
locality named Cozzo di Mastro Pasquale (Fig. 1). The
systematic-taxonomic study has allowed some considerations about belemnite biostratigraphy and palaeobiogeography. A fauna was first described from this outcrop
in a preliminary study by Combémorel et al. (1994a).
Continued sampling over the last fifteen years produced
the collection described in this paper, representing a faunal assemblage that is unique in Italy, but is well known
from coeval beds in Romania, Bulgaria, France and
southern Germany as well as the Caucasus.
The European and Mediterranean Aalenian-Bajocian
belemnites are poorly studied either for the lack of outcrops or for the limited number of specimens. The only
Fig. 1 - Location map of the examined outcrop (*).
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authors who report some of the taxa analyzed here are
Schwegler (1938, 1965, 1971) and Riegraf (1980, 1981),
who discuss the stratigraphic and geographic distribution of the Jurassic belemnite genera Megateuthis,
Mesoteuthis, Eocylindroteuthis, Homaloteuthis, Holcobelus, Brevibelus, Pachybelemnopsis and Hibolithes.
Schlegelmilch (1998) figures some original specimens
Schwegler (1938, 1965, 1971) that were only known
after drawings. Althoff (1928) and Waldeck (1975)
record Megateuthis from the Bajocian of Bielefeld and
Lower Saxony (Germany). Kumm (1952) cites
Homaloteuthis and Megateuthis from the Upper
Aalenian and Lower Bajocian of Lower Saxony. Buser
(1952) and Lieb & Bodmer (1955) record Megateuthis,
Holcobelus and Brevibelus from the Swiss Jurassic. V.
Hochstetter (1897) reports Lower Bajocian belemnites
from St. Veit near Vienna (Austria), including
Brevibelus, Megateuthis, Pachybelemnopsis and a new
species, Belemnites eduardi. Stolley (1927) includes the
latter into his new genus Holcobelus. Phillips (1868) discusses species of Megateuthis, Holcobelus and
Pachybelemnopsis, but unfortunately no recent, more
detailed studies on these forms are presently available.
From Belgium, Luxembourg and NE France (AlsaceLorraine) Chapuis & Dewalque (1854) describe for the
first time Bajocian belemnites (Megateuthis), while
Lepsius (1875), Branco (1879), Bleicher (1884), Haug
(1886), Van Werveke (1901), Benecke (1905, 1905),
Laux (1921-23), Lucius (1945, 1948) and Maubeuge
(1955) report several specimens referable to
Homaloteuthis, Brevibelus, Megateuthis and Pachybelemnopsis. Weis (2006) describes a Lower Bajocian
(Humphriesianum Zone) Megateuthis-Pachybelemnopsis assemblage from Luxembourg. The belemnite
monograph by Eudes-Deslongchamps (1878) is remarkable for NW France (Normandy). The author describes
Megateuthis, Brevibelus and Pachybelemnopsis together
with several new Upper Aalenian species; unfortunately
it has not been revised up to now. For these latter forms
Stolley (1927) institutes the new genus Holcobelus.
Riche (1904), Lissajous (1925) and Roché (1939)
record Holcobelus, Brevibelus, Megateuthis and
Pachybelemnopsis from Central France (Burgundy),
while Garnier (1872), Zürcher (1891), Roman &
Gennevaux (1912), Gérard (1936) and Riegraf (1980)
report Holcobelus and Pachybelemnopsis from the
Aalenian/Bajocian boundary of Southern France.
In SE Europe the distribution pattern appears to be
somewhat more complete. Stoyanova-Vergilova (1982,
1985, 1990, 1993) gives an extensive overview of
Aalenian and Bajocian belemnites from Bulgaria. The
author quotes and figures several species of Holcobelus,
Brevibelus, Megateuthis, Paramegateuthis, and Pachybelemnopsis. A comparable assemblage with Megateuthis, Brevibelus, Holcobelus, Pachybelemnopsis and
Hibolithes is reported by Preda (1975) from the Bajocian
of Romania. Such faunas are also reported from
Caucasus and Crimea (Krimholz 1947, 1953). In
Southern Italy, as mentioned above, Combémorel et al.
MARIOTTI et al.
(1994a) report an isolated belemnite assemblage of
Aalenian-Early Bajocian age from Caloveto - the locality which is the object of the present study - with
Holcobelus, Brevibelus, Pachybelemnopsis, Hibolithes
and Megateuthis.
GEOLOGICAL SETTING
The backbone of the southernmost region of peninsular Italy, Calabria, consists of Early Palaeozoic metamorphites which are intruded by Late Palaeozoic
(Hercynian) granitoids (Amodio-Morelli et al., 1979,
and bibliography therein; Dubois,1976; Bouillin, 1984).
In the area located at the north-eastern, Ionian, side of
Calabria, named Sila Greca, the Palaeozoic crystalline
basement is transgressively covered by MesozoicPalaeogene sediments. This complex was then subjected
to Alpine/Apenninic orogenic deformation, and followed
by a new Miocene-Quaternary sedimentary cycle.
The Mesozoic unmetamorphosed sedimentary cover is
characterized by two distinct successions: the Longobucco Group and the Caloveto Group, differing in their
facies, thickness, and chronostratigraphic range (Teale,
1985; Young et al., 1986, and bibliography therein;
Santantonio & Teale, 1987; Teale & Young, 1987;
Bouillin et al., 1988) (Fig. 2).
The paper by Young et al. (1986) contains a useful list
of references - to which the reader is referred - documenting the early efforts of palaeontologists, mostly
ammonite specialists, who tackled the stratigraphy of the
area around the turn of the nineteenth century.
Importantly, the papers by Young et al. (1986) and
Santantonio & Teale (1987) also introduce a new - as yet
informal - lithostratigraphy, which will be used here.
The Longobucco Group
The Longobucco Group (Triassic-Jurassic boundary to
Toarcian) (Fig. 2) is exposed across an area of about 150
km and it is ~1.5 km thick. It documents sedimentation
on a rifted continental margin, in environments evolving
from continental (fluvial deposits), to carbonate shelf, to
slope and finally to a deep turbiditic basin (see also
Zuffa, 1980).
The Longobucco Group is composed by the following
units:
1) - Torrente Duno formation (Rhaetian-Hettangian):
sandstones (mainly quartzarenites), shales and conglomerates (red beds) of fluvial environment, 100-200 m
thick;
2) - Bocchigliero formation (Sinemurian s.l.): blackish
limestones (wackestones to grainstones), mostly oolitic,
with a rich shelly fauna (brachiopods and bivalves), also
with occasional corals, oncoids and Thalassinoides burrows. Material derived from the crystalline basement
(e.g. lenses of cross bedded quartz conglomerates) and
plant remains (e.g. tree logs) are abundant throughout.
Thickness up to about 100 m;
AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE ...
3) - Fosso Petrone formation (Lower Pliensbachian):
bioturbated marls with bivalves, belemnites, ammonites
and plant remains; about 200 m thick; shelf-edge or
slope facies;
4) - Fiume Trionto formation (Upper PliensbachianLower Toarcian): mixed siliciclastic/carbonate turbidites
and interbedded hemipelagites, bearing olistoliths, tens
to a few hundred metres across, made of the first two
units of the Group. Up to ~1200 m thick.
This succession bears the evidence that the evolution
from a continental fluvial environment to a deep marine
environment must have been driven by rift tectonics in
the Early Jurassic (Santantonio, 1993), ultimately providing the accommodation space for the thick, high sedimentation-rate Trionto formation turbidites.
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Submarine tectonic escarpments exposing the local
shelf stratigraphy were in this view the source areas of
the megaclastic rockfall and debris flow material, and
the turbidites are indeed seen to onlap the Palaeozoic
phyllites in several basin-margin localities across the
Bocchigliero area. The continental deposits (Torrente
Duno formation) gradually evolve to intertidal and to
subtidal marine deposits (Bocchigliero formation).
Facies range from quartzose conglomerates and cross
bedded hybrid arenites to marls, to cross laminated
oolitic grainstones and bioturbated subtidal wackestones
bearing a diverse marine shelly fauna. The Bocchigliero
formation documents the development of a mixed carbonate/siliciclastic shelf hosting oolite bars, under the
influence of river deltas providing abundant siliciclastic
input and plant material sourced by a crystalline hinterland subjected to a humid climate, and then reworked by
tidal and alongshore currents. The transition to the next
unit is documented by subtidal carbonate mud facies
with neritic fauna and flora and the gradual disappearance of pebble-and sand-grade siliciclastic material,
indicating distal open shelf conditions with decreasing
ambient energy. Onset of the Petrone formation is
marked by the appearance of cephalopods (belemnites
and ammonites) and the occurrence of thin fine-grained
turbidites, an indication of a deeper-water shelf to slope
environment (Young et al., 1986). The lower Toarcian
top of the Trionto formation turbidites is cut by modern
erosion, so the later Jurassic sedimentary evolution of
the Longobucco Group is unknown.
The Caloveto Group
Fig. 2 - Correlated sequences of the Longobucco and Caloveto Groups.
Longobucco Group: 1- Torrente Duno Formation; 2- Bocchigliero
Formation; 3- Fosso Petrone Formation; 4- Trionto Formation; 5Hercynian basement (metamorphites); 6- Hercynian basement (granitoid). Caloveto Group: 1- Lower Caloveto Formation; 2- Upper
Caloveto Formation; 3- S.Onofrio Formation (basal red marls); 4S.Onofrio Formation (pelagic limestone, radiolarites with admixed
siliciclastic and calciclastic turbidites); 5- Hercynian basement (metamorphites); 6- Hercynian basement (granitoids). Dashed lines are time
lines.
The Caloveto Group (Fig. 2) represents a sequence
sedimented around and on a basement structural high,
and as such spans a long interval of geological time
(Early Jurassic to Early Cretaceous) through a much
reduced thickness (maximum 300 m). The Longobucco
Group acted in the Early Jurassic as a strongly subsiding
sediment trap separating the offshore highs of Caloveto
from the Jurassic mainland Calabria. The Caloveto
Group is exposed in small, isolated outcrops dissected
by faults, which often makes bed correlation difficult.
The main outcrops of the Caloveto Group are found in
two main areas: 1. nearby the village of Caloveto, and 2.
along the Colognati stream valley. The latter area is the
sole having exposures of the younger (Upper Jurassic
and Neocomian) part of the succession. The deposits of
the Caloveto Group display a complex array of unconformable contacts and strongly variable geometries and
facies, documenting sedimentation on a rugged substrate, under a very strict sinsedimentary tectonic control.
The Caloveto Group is composed by the following
units, from the base:
1) - Lower Caloveto formation (Pliensbachian). This
carbonate unit marks a marine transgression over an
intrusive and metamorphic Hercynian basement. It is
made of up to ~80 m of pale-coloured lime grainstones
4
and packstones and hybrid arenites, cross-bedded in
places, bearing rich shallow water benthic faunas (gastropods, bivalves, crinoids, corals, Tubiphytes, etc.).
Low-energy cyclic mud-rich facies are remarkably
missing in this unit. This formation also bears an up to
40 m thick conglomerate facies, with clast- to matrix
(coarse carbonate sand)- supported conglomerates bearing rounded pebbles derived from the basement. These
conglomerates are diachronous, marking the irregular,
non conformable contact to the phyllites at Caloveto
(more on this below). The change to the next unit can be
either sharp, through a mineralized paraconformity, or
transitional, and made of an up to 10m thick package of
pink brachiopod- (mostly rhynchonellids) to crinoid-rich
packstones. The age of the unit is poorly constrained, but
the top levels bear Agerina martana (Farinacci), a
Pliensbachian benthic foram.
2) - Upper Caloveto formation (Toarcian, Serpentinus
to Dispansum Zones), about 0.5-8 m of red pelagic,
whole-fossil lime wackstones and nodular marly mudstone (ammonitico rosso facies). These red pelagites
often fill deep fractures which cross the former unit as
well as the basement in the form of neptunian dykes.
Ammonites, echinoderms (mostly crinoids), brachiopods and posidoniid bivalves characterize this unit.
A sedimentary hiatus of some 2 My must exist at the
Pliensbachian-Toarcian boundary, corresponding to the
iron-stained basal surface. The ammonite assemblage
recovered indicates the Lower (not lowermost) Toarcian
Serpentinus Zone.
3) - Sant’Onofrio formation (Late Toarcian to Hauterivian), maximum thickness of 150 m, comprising red
bioturbated (Zoophychos) (Fig. 3) marls, Aptychus limestone, radiolarian cherts and calpionellid limestone.
These “background” lithologies host a variety of spectacular gravity flow deposits (rock falls, debris flows,
turbidites) in the outcrop areas of the high Colognati
Fig. 3 - Zoophychos from red marls (lower part of the Sant’Onorio
Formation).
MARIOTTI et al.
stream valley, near Rossano (about 15 km west of the
Caloveto outcrop area). These deposits include widespread hybrid arenites, clasts of the crystalline basement
up to boulder size, and peloid-oolite calcarenites from
unknown carbonate platform source areas. The various
lithologies are, at present, dealt with as informal members, but may end up being treated as formations, should
the Sant’Onofrio be elevated at group rank. Ammonites,
belemnites and posidoniid bivalves occur in the red
marls. Apart from aptychi, ammonites are virtually
absent in the post-Bajocian sediments, in which
belemites and other fauna occur sporadically.
The Caloveto Group indicates the initial development
of carbonate aprons around islands made of Palaeozoic
rock. These carbonates have great lateral variability, but
their ubiquitous grainstone to conglomerate facies is an
evidence for high energy conditions in a coastal environment. Coral patches occurred sporadically, but nowhere
did an inner lagoon with a low-energy muddy facies
exist. By contrast to the Longobucco basin, that initially
developed as a wide shelf backed by a continent with a
fluvial system, the Lower Caloveto formation was sedimented due to subsidence of emerged land offshore, and
these islands were too small to host a proper drainage
system. Thence the white (as opposed to black) color of
the limestone, and the lack of any plant material. These
carbonates are locally seen to fill fractures in the phyllite
basement, suggesting that subsidence had a strong tectonic component.
Synsedimentary faulting was certainly most active in
the Toarcian, and this probably had two main effects: 1.
the foundering and drowning of any shallow water carbonate system, and 2. the tectonic dissection of both the
basement and the Lower Caloveto limestone. This produced a network of neptunian dykes, and set the conditions for the development of various unconformable contacts, due to the birth of a markedly irregular submarine
topography. The Colognati area has megaclastic beds
sealed by late Toarcian ammonite-rich deposits
(Santantonio & Teale, 1987; Santantonio, 1993), so
faulting was active across both the main outcrop areas
mentioned above.
The deposits of the Sant’Onofrio formation exhibit
various types of geometric relationships with their substrate. In the Caloveto area contacts vary from paraconformities to angular unconformities, both with the basement and with the faulted shallow-water limestone. At Il
Torno (Colognati Valley), Aalenian marls with
Zoophychos are seen to drape the Toarcian megabreccia
as well as to onlap the granite basement.
The youngest beds seen at Caloveto are the radiolarian
chert facies (?Callovian-?Oxfordian), which, where resting unconformably on the Lower Caloveto formation, are
seen to induce the growth of chert nodules in the latter,
by analogy with basin-margin unconformities in the
Apennines (Santantonio et al., 1996; Galluzzo &
Santantonio, 2002). The Aptychus limestone is found in
the Colognati area, and is seen to change into a calpionel-
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Fig. 4 - Panoramic view of the Cozzo di Mastro Pasquale outcrop near Caloveto. Dashed line is the trace of the basal unconformity of onlapping
marls (lower part of the Sant’Onofrio Formation).
lid limestone (Maiolica facies) which is markedly lithoclastic (basement clasts, plus clasts of the older formations) and also bears turbiditic calcarenites. These features might be indicative of a reactivation of Early
Jurassic faults around the Tithonian/Berriasian boundary.
The “Cozzo di Mastro Pasquale” outcrop
At km 8.2 along the road connecting the villages of
Caloveto and Bocchigliero, in a locality named “Cozzo
di Mastro Pasquale” (Fig. 1), it is possible to observe: a)
the top of the Lower Caloveto formation; b) the base of
the red nodular limestone of the Upper Caloveto formation; c) the marls of the Sant’Onofrio formation.
a) The top of the Lower Caloveto formation is an erosional surface, locally iron-stained. One remarkable outcrop at a narrow topographic terrace demonstrates the
richly fossiliferous, condensed, nature of the last bed of
the unit. This bears ostreids, echinoids, gastropods, brachiopods, and set in a matrix of hybrid grainstone (crystalline granules). The first occurrence of rare ammonites
(Protogrammoceras sp., Pliensbachian) indicates the
onset of a pelagic influence.
b) This area also features the best (though not the
thickest) outcrop of the Upper Caloveto formation. This
is made of red pelagic limestone and marly limestone,
with echinoids in the lower part, brachiopods and
ammonites of Early Toarcian age (Serpentinus Zone).
The first centimetres of the Upper Caloveto formation
are a red condensed wackestone bearing specimens
ascribed to Hildaites serpentinus. Marly limestone with
Hildoceras sublevisoni and Mercaticeras sp. (middle
Toarcian) follows, attaining a maximum thickness of 1
m. These levels show rare crystalline clasts.
c) The marls of the Sant’Onofrio formation (Fig. 4)
that follow have also a variable thickness (few metres to
>25m), due to their unconformable base. They are made
of thick bioturbated beds alternating with thin laminated
posidoniid-rich beds. Soft marl clasts with a thin ironrich coating could be the product of reworking of incipient hardgrounds by burrowers. About 7 m above the top
of the Lower Caloveto limestone in the section described
earlier for the Toarcian, which displays about 16 m of
Sant’Onofrio marls, a level with large (10-15 cm longer
axis) bivalves probably belonging to the genus
Inoceramus also produced a rich Aalenian ammonite
fauna. This is composed by Tmetoceras scissum
(Benecke, 1865), Ancolioceras opalinoides (Mayer),
Erycites fallifax Arkell 1957, Erycites intermedius
(Hantken in Prinz, 1904) (Fig. 5) and lytoceratids found
in situ from 6 m to 11 m above the stratigraphic base of
the unit. Tmetoceras scissum indicates the early
Aalenian (Opalinum Zone; Venturi 1982; Venturi &
Ferri, 2001; Pallini et al., 2004). Erycites intermedius
appears at the base of the Opalinum Zone (Early
Aaleniano), while Erycites fallifax is generically an
Aalenian form. Ancolioceras opalinoides (Mayer) is a
marker of the Opalinoides Subzone of the Murchisone
Zone. This assemblage therefore covers the middle part
of the Opalinum Zone to the middle part of Murchisonae
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MARIOTTI et al.
Fig. 5 - a) Erycites intermedius, Early Aalenian, base of the Opalinum Zone, x 0.75 ; b-c) “Docidoceras” cf. perfectum, Discites Zone, x 0.75; d
Tmetoceras scissum, Aalenian, Opalinum Zone, x 1.
Zone (Cresta, 2002; ed.). Two stephanoceratids were
found in the upper part of red marls, one in a displaced
block, the other in place. They belong to Docidoceras,
probably a “D.” cf. perfectum (Fig. 5) (see Cresta &
Galacz, 1990; Pl. II, fig. 4), which indicates the Early
Bajocian Discites Zone in the Umbria-Marche Apennines. The same species is reported by Callomon &
Chandler (1990) from beds of Late Aalenian-Early
Bajocian age in Dorset (Southern England). In exposures
other than those of the Cozzo di Mastro Pasquale area,
the lower part of the marl succession has also yielded
ammonites indicative of a Late Toarcian (Aalensis Zone)
and Early Aalenian (Opalinum Zone) age. The presentday topography, and an unconformable contact with
Miocene conglomerates and sandstone, truncate the
upper part of the marls, which are seen to change conformably to red radiolarites a couple of metres above the
Docidoceras-bearing bed.
The belemnites studied in this paper were collected
from the red marls of the Sant’Onofrio Formation (Fig.
6), and their stratigraphic distribution (Aalenian-Early
Bajocian; Combémorel et al., 1994a) is in good agreement with ammonite stratigraphy as established in the
area.
An age for the top of the Sant’Onofrio Formation can
be tentatively placed in the Bajocian.
MATERIAL AND METHODS
The terminology herein used is derived from Doyle &
Kelly (1988) and Mariotti (2003). Individual descriptions also include size measurements in the cases of
complete or almost complete specimens. All measures
are in millimeters. Estimated values for incomplete specimens are marked by an asterisk (*). Dimensional adjectives are used as in Doyle & Kelly (1988); the terms
small, medium and large, related to the length of the rostrum (L), are respectively referred to L < 80 mm, L
between 80 and 110 mm and L > 110 mm.
Abbreviations as in Mariotti (2003): L, total preserved
length; Dv, dorso-ventral diameter at alveolar opening;
Dl, lateral diameter at alveolar opening; Dvmax, maximum ventral diameter; Dlmax, maximum lateral diameter; X, length from apex to Dmax; Ic, compression index,
the ratio between dorsal-ventral diameter and lateral
diameter, calculated at the level of the alveolar opening
(Ica), and at the level of the maximum dorso-ventral
diameter (Icm); Id, depression index, the ratio between
lateral diameter and dorso-ventral diameter calculated at
the level of the alveolar opening (Ida), and at the level of
the maximum lateral diameter (Idm).
All the collected specimens are stored in the
Palaeontological Museum of the Dipartimento di
Scienze della Terra of the University “La Sapienza” of
Rome under collection number NS 20/…
The rostra collected are not in a good state of preservation, even if often they are complete. The phragmocone is never preserved but frequently the alveolus is
observable. Some of the rostra as well as the ammonites,
preserved always as inner moulds, show strong tectonic
stress deformation (Fig. 7 a-c).
Furthermore, some specimens exhibit borings made by
acrotoracic cirripeds (Fig. 7 d). These organisms are
marine sessile crustaceans with a boring habitat in carbonate substrata. They form in the substratum a sacshaped cavity with a comma-shaped opening (with a
round end and a sharp end), with a total length only
rarely exceeding 2 mm. These cirripeds orient the cirri
against the current flux from the rounded portion of the
opening directing them towards the sharpened portion
(Fig. 8) (Petriconi, 1971). Assuming that borings postdate death of the belemnites and settlement of the rostrum on the sea-bottom, it is conceivable that sedimenta-
7
Fig. 8 - The boring of an acrotoracic cirriped (after Petriconi, 1971).
SYSTEMATICS
The classification adopted here is based on Riegraf
(1995) and Riegraf et al. (1998).
Order Belemnitida MacGillivray, 1840
Suborder Belemnitina MacGillivray, 1840
Family Megateuthididae Sachs & Nalnjaeva, 1967
Fig. 6 - Belemnites bearing red marls of the lower part of the
Sant’Onofrio Formation. The white arrow indicates a hammer.
tion rates were as slow as to permit the colonisation of
rostra before they were covered by sediment. Sometimes
the borings reach the innermost part of the rostrum (Fig.
7 d).
Genus Brevibelus Doyle, 1992
Diagnosis
“Small, short and robust, cylindriconical to conical
Megateuthididae. Outline symmetrical and conical to
cylindriconical, profile symmetrical to slightly asymmetrical, otherwise similar to the outline. Apex obtuse to
moderately acute, often mucronate. Venter inflated in
some species. Transverse sections quadrate, compressed
in some species, depressed in others. Apex devoid of
grooves or striae. Lateral lines may be well-developed,
consisting of two weak and parallel depressions separated by a relatively well-developed ridge. The phragmocone is ventrally displaced, penetrating one half to one
third of the rostrum. Apical lines strongly cyrtolineate,
apical angle approximately 27°.” (After Doyle, 1992)
Type species
Belemnites breviformis Voltz, 1830
Remarks
Parapassaloteuthis Riegraf, 1980 is the only genus similar to Brevibelus, but the latter differs in having a more
conical rostrum, no apical groove and weaker lateral
lines.
Brevibelus breviformis (Voltz, 1830)
Fig. 7 - a-c) boudinaged rostra, x 1; d) borings of acrotoracic cirripedes
at the outer surface (left) and in the innermost part of the guard, x 1.
(Pl. 1, Fig.1 a-c)
8
MARIOTTI et al.
*1830 Belemnites breviformis Voltz, p. 42, Pl. II, Figs. 24.
1992 Brevibelus breviformis - Doyle, 62, Pl. 23, figs. 6,
10, 11; Pl. 24, figs. 1, 2. (cum syn.).
1999 Brevibelus breviformis - Weis, 222, Figs. 4-5, 2729
Type species
Belemnites giganteus v. Schlotheim, 1820 (= Belemnites
Aalensis Voltz, 1830) by subsequent designation
(Douvillé, 1879, p. 91)
Material
Two complete juvenile specimens in mediocre state of
preservation (NS 20/877-/1184).
(Pl. 1; Fig. 12 a-c)
Description
The small rostrum is typically conical and slender. The
outline is symmetrical and conical, the profile is nearly
symmetrical with a slightly flattened venter. The apex is
moderately acute. The transverse section is subquadrate
in the alveolar region and becomes elliptical in the apical region. No groove is present. The phragmocone penetrates one half of the rostrum.
Dimensions
See Tab. 1
Remarks
The characteristic conical and slender shape and the lack
of groove permit a systematic attribution to a juvenile
form of Brevibelus. The juvenile stadium of the two
specimens is based on ontogenetic observations of Weis
(1999, p. 224). The only similar species is Brevibelus
gingensis, but Brevibelus breviformis is different for its
more slender shape.
In Italy this species, common in the European belemnite
assemblages, is recorded only from the “Cozzo di
Mastro Pasquale” outcrop.
Geographical and stratigraphical distribution
Upper Toarcian to Lower Bajocian of Europe, Russia,
western Canada (Doyle, 1992), and Aalenian-Lower
Bajocian of southern Italy.
Megateuthis sp.
Material
One juvenile fragment showing the alveolar and partially
the stem region (NS 20/819).
Description
The small conical rostrum is characterized by two apical
dorso-lateral grooves visible on the transverse section. It
lacks a ventral groove and is strongly compressed with
an elliptical cross section. The alveolus probably penetrates one fourth of the rostrum.
Remarks
The fragmentary preservation does not permit a specific
attribution. The blunted apical part of the rostrum exposes the typical juvenile stages of Megateuthis with clear
indications of two apical dorso-lateral grooves. Both
Schwegler (1965) and Pugacewska (1961) illustrate similarly short, compressed and conical juveniles of
Megateuthis elliptica from the Bajocian of Poland and
southern Germany. The fragmentary state (lack of apical
region) of the unique collected specimen does not allow
drawing any further comparison.
Bajocian of Europe, ?North America (Doyle, 1992), and
Aalenian-Lower Bajocian of southern Italy.
Suborder Pachybelemnopseina Riegraf, 1998
Family Holcobelidae Gustomesov, 1977
Genus Holcobelus Stolley, 1927
Diagnosis
Small to medium sized, elongated, conical to cylindrical
Holcobelidae. Apex acute or obtuse (orthorostrum). No
apical grooves. Intermediate ventral groove extending
from the alveolar region to nearly the apex. Transverse
sections rounded or elliptical, compressed. Epirostrum
well-developed in some species.
Genus Megateuthis Bayle, 1878
Diagnosis
Very large conical to elongate cylindriconical Megateuthididae, with a well-developed epirostrum. The outline
and profile are symmetrical and conical to cylindriconical. The apex is obtuse and bears dorso-lateral grooves
and generally well-marked striae. Transverse sections
are compressed and elliptical.
Remarks
The phylogenetic affinities of Holcobelus were
discussed in Combémorel et al. (1994a).
Type species
Belemnites munieri Eudes-Deslongchamps, 1878.
Holcobelus trauthi Stolley, 1927
(Pl. 1, fig. 3 a-c)
9
*1927 Holcobelus trauthi Stolley, p.
118, pl. XXIV, fig. 7-8.
1994 Holcobelus trauthi - Combémorel
et al., p. 47, Pl. 1, fig. 4.
Material
Six complete rostra (NS 20/1181,
/1179, /862, /1178, /1400, /1403), one
juvenile form (NS 20/828), one specimen lacking the alveolar region (NS
20/815), and several fragments of the
distal part of the alveolar region (NS
20/786, /823, /776, /866).
Description
The middle sized rostrum is conical in
the posterior. Outline and profile are
symmetrical and cylindriconical, very
slightly hastate in the adult. The ventral groove is narrow
and well-marked, reducing towards the apex which it
appears to meet. The transverse section is sub-rectangular and compressed. The apical region is slightly obtuse.
Dimensions
See Tab. 3
Dimensions
See Tab. 2
Remarks
This species is similar to the original specimens of
Stolley (1927, pl. XXIV, fig. 7-8). The closest species is
Holcobelus subblainvillei but the latter shows a more
conical slender shape and less compression.
Lower Bajocian of Normandy (France), and AalenianLower Bajocian of southern Italy.
Holcobelus subblainvillei (Eudes-Deslongchamps, 1878)
(Pl. 1, Fig. 7 a-c)
*1878 Belemnites subblainvillei Eudes-Deslongchamps,
p. 60, pl. V, figs. 15, 17 , pl. VII, figs. 5-9 (cum syn.)
1927 Holcobelus subblainvillei - Stolley, p. 118, pl.
XXIV, fig. 5.
Material
Five complete rostra (NS 20/897, /1404, /785, /784,
/880) and several fragments.
Description
The conical rostrum is medium-sized and slender. The
profile is symmetrical and conical; the outline is
symmetrical, but less conical than profile. A ventral
narrow deep groove extends from the alveolus next to
the apex. The apical region ends with an acute and
slightly mucronate point. The transverse section is
subcircular to slightly compressed. The phragmocone
penetrates one third of the rostrum.
Remarks
This species is very similar to Holcobelus blainvillii, but
it differs by its groove extending from alveolus to apex.
Another taxon comparable with Holcobelus subblainvillei is Holcobelus munieri. Holcobelus munieri has a
very long and more slender rostrum and an apical region
more acute.
Combémorel et al. (1994b), in the revision of
d’Orbigny’s collection, figured a longitudinal section of
Holcobelus munieri with an epirostrum, and retained
that Holcobelus subblainvillei would be Holcobelus
munieri without epirostrum. This might be truth, but for
the moment we do not have appropriate material for further investigation.
Upper Aalenian and Lower Bajocian of Normandy,
Haute-Provence (France), and Aalenian-Lower Bajocian
of southern Italy.
Holcobelus munieri (Eudes-Deslongchamps, 1878)
(Pl. 1, Figs. 2 a-c, 5 a-c)
*1878 Belemnites munieri Eudes-Deslongchamps, p. 63,
Pl. V, figs. 3-6, 12-14; Pl. VI, figs. 5-11. (Cum syn.)
1993 Holcobelus munieri Stoyanova-Vergilova, p. 73,
Pl. XXXV, figs. 1-4; Pl. XXXVI, figs. 1-4. (Cum syn.)
10
Material
Two juveniles (NS 20/880, /843), two specimens lacking
the distal end (NS 20/838, /768) and one specimen
lacking the most part of the alveolar region (NS 20/878).
Description
Very slender conical and slightly compressed rostrum.
The outline and profile are symmetrical and conical. The
dorsal, the ventral sides and the flanks converge weakly
but continuously till the apex. The apical region is acute
and the end is sometimes very slightly obtuse. A deep
ventral groove starts from the alveolar opening, persists
into the stem region disappearing at the apex. The transversal section is weakly compressed, subquadrate. The
phragmocone penetrates one fourth of the rostrum. The
adult stage presents an epirostrum.
MARIOTTI et al.
adult specimens. The transversal section is compressed
and subrectangular. A deep ventral groove extends from
the alveolus till the apex. The phragmocone, slightly
eccentric ventrally, penetrates for one third of the total
length of the rostrum.
Dimensions
See Tab. 5
Dimensions
See Tab. 4
Remarks
Remarks
The small-sized rostrum, the compressed subrectangular
transversal section and the obtuse apical region make
this species unique within the genus Holcobelus.
Upper Aalenian of Normandy, Haute-Provence (France)
and Aalenian-Lower Bajocian of southern Italy.
Holcobelus tschegemensis (Krimholz, 1931)
This species differs from the other species ascribed to
Holcobelus by the long slender conical rostrum, the
conical apical region and the long groove.
Upper Aalenian and Lower Bajocian of Normandy,
southern France, Bulgaria, and Aalenian-Lower Bajocian of southern Italy.
Holcobelus tetramerus (Eudes-Deslongchamps, 1878)
(Pl. 1, Figs. 4 a-c, 6 a-c; 8 a-c)
*1878 Belemnites tetramerus Eudes-Deslongchamps, p.
67, Pl. VII, figs. 10-20.
1927 Holcobelus tetramerus - Stolley, p. 118.
Material
Five complete specimens (NS 20/774, /773, /892, /1399,
/1405).
Description
The rostrum is small-sized and compressed. The outline
is symmetrical and conical, the profile is symmetrical
and cylindriconical. The flanks and the ventral and lateral sides run parallel to converge suddenly in the apical
region ending with an obtuse point; this is evident in the
(Pl. 1, Fig. 11 a-c)
*1931 Belemnopsis tschegemensis Krimholz, p. 27, pl. I,
figs. 26-32.
1990 Holcobelus tschegemensis - Stoyanova-Vergilova,
pl. I, fig. 3.
1993 Holcobelus tschegemensis - Stoyanova-Vergilova,
pl. XXXVII, fig. 4.
Material
Two juvenile specimens (NS 20/824, NS 20/851), one
specimen lacking the distal end (NS 20/775), one
specimen lacking the apical region (NS 20/1406), three
fragments of the apical region (NS 20/864, /907, /1412),
one rostrum lacking the terminal end and part of the
alveolar region (NS 20/865) and two specimens without
the apical region (NS 20/847, /777).
Description
The medium-sized rostrum is typically slender and thin.
The profile and outline are symmetrical and cylindrical.
The transverse section is elliptical to subquadrate and
weakly compressed for all the length of rostrum. The
alveolus penetrates one sixth of the rostrum. The long
ventral groove is deep in the anterior region, and fades
slightly out towards the apex.
Dimensions
See Tab. 6
11
Material
Three juvenile specimens (NS 20/788, /789, /1401) all
without phragmocone and alveolus.
Description
Small-sized fusiform rostrum with outline and profile
symmetrical and fusiform. The maximum width of rostrum occurs between 1/3 and 1/4 from the apex. A very
thin and narrow ventral canal is present in the most proximal part. Weak lateral lines are visible on the flanks.
The transverse section is more or less circular.
Remarks
This species, characterized by a medium-sized slender
an thin rostrum and a long ventral groove, is outstanding
in the genus Holcobelus.
Dimensions
See Tab. 7
Upper Aalenian of the Caucasus, Bulgaria, Haute
Provence (France) and Aalenian-Lower Bajocian of
southern Italy.
Family Mesohibolitidae Nerodenko, 1983
Genus Hibolithes Denys de Montfort, 1808
Diagnosis
Elongated, slender Mesohibolitidae. Both profile and
outline symmetrical and usually hastate. Maximum
transverse diameter usually at the apical region.
Transverse section generally circular, in some cases
depressed in the apical region in some others compressed in the alveolar region. Shallow ventral canal limited at the anterior half of the rostrum. Well evident lateral lines. Central apical line.
Type species
Hibolithes hastatus Denys de Montfort, 1808
Remarks
The two juvenile specimens are ascribed to Hibolithes
wuerttembergicus by their fusiform shape, the weak
alveolar canal and lateral lines. They are synonymous
with the juveniles figured by v. Zieten (1830-33, Pl. 25,
fig. 3c-f). Pugaczewska (1961) describes and figures
three ontogenetic stages for this species: nepionic, neanic and ephebic-gerontic. The here analyzed specimens
could be nepionic forms for their fusiform shape, the
length and the tapering of the terminal portion and the
poorly incised alveolar canal.
Stratigraphical range and geographical distribution
Bajocian to Bathonian of Germany, Luxembourg,
Poland and Aalenian-Lower Bajocian of southern Italy.
Hibolithes wuerttembergicus (Oppel, 1856)
Hibolithes sp.
(Pl. 1, Fig. 10 a-c; Pl. 2, Figs. 4 a-c, 8 a-c)
(Pl. 2, Fig. 5 a-c)
*1856 Belemnites württembergicus Oppel, p. 485.
1830 Actinocamax cf. milleri - v. Zieten, p. 33, Pl. 25,
fig. 3a.
1830 Actinocamax fusiformis - v. Zieten, p. 33, Pl. 25,
fig. 3b.
1832 Actinocamax lanceolatus Hartmann - v. Zieten,
p.33, Pl. 25, figs. 3c-f.
1961 Hibolithes wuerttembergicus - Pugaczewska, p.
177, fig. 23, Pl. XXI, fig. 1-8.
1981 Hibolithes (H.) wuerttembergicus - Riegraf, p. 6770, figs. 37-38 (cum syn.).
1998 Hibolithes wuerttembergicus - Schlegelmilch,
p.82, Pl. 18, figs. 6-9, Pl. 20, fig.14.
v. 2005 Hibolithes wuerttembergicus - Weis & Gross,
p.68, fig. A3.
v. 2006 Hibolithes wuerttembergicus - Weis, p. 161, fig.
11.
Material
Three stem fragments close to the alveolar region (NS
20/779, /1395, 848) and one fragment showing the
alveolar region and partially the stem region (/1413).
Description
Fragments of medium and large rostra with a probable
fusiform shape. A clear broad ventral canal is observable. The dorsal side shows a short depression extending
distally as a flattened area. The transverse sections are
circular to slightly elliptical. Two lateral lines are present
on each flank.
Remarks
The type of the canal, the transverse section, the presence of lateral lines and a short dorsal depression permit
12
to ascribe these fragments to Hibolithes.
Genus Pachybelemnopsis Riegraf, 1980
Diagnosis
Cylindrical to hastate, elongate Mesohibolithidae.
Outline symmetrical. Profile asymmetrical. Apex acute.
Transverse section generally depressed in the apical and
stem regions. Broad, ventral alveolar canal sometimes
extending up to the apex. Apical line ventrally eccentric.
Lateral lines rarely present.
Type species
Belemnites canaliculatus v. Schlotheim, 1820
Pachybelemnopsis baculiformis Riegraf, 1980
(Pl. 1, fig. 13 a-c, Fig. 9 a-c; Pl.2, Figs. 7 a-c, 9 a-c)
*v.1980 Belemnopsis (Pachybelemnopsis) baculiformis
Riegraf, p. 179-181, text-figs. 160-161, Pl. 1, fig. 11
[cum syn.]
1998 Belemnopsis baculiformis - Schlegelmilch, p. 79,
Pl. 16, figs. 10-11
v. 2006 Pachybelemnopsis baculiformis - Weis, p. 161,
fig. 12
Material
One juvenile specimen (NS 20/830), three specimens
lacking the alveolar region (NS 20/837, /850, /834), and
two fragments of the stem region (NS 20/839, /822).
Description
Medium sized rostrum with a fusiform shape narrowing
distally to end by a mucronate tip. The transverse section
is subcircular in the alveolar region and more or less
depressed towards the apex. A deep ventral canal
extends towards the point, it is broad with clear ridges
and its thickness decreases distally; it fades out at the
beginning of the apical region. The lateral lines are
weak. The outline is hastate, with maximum diameter
located at the 2/3 of the total preserved length. The
profile is hastate, slightly asymmetrical. The apical
region is short and slightly obtuse.
Dimensions
See Tab. 8
MARIOTTI et al.
Remarks
Pachybelemnopsis baculiformis distinguishes from other
species of Pachybelemnopsis by its little club-like form
and the relatively long and broad ventral canal.
Bajocian (Humpresianum Zone) south-western Germany,
north-eastern France, Luxembourg and Aalenian-Lower
Bajocian of southern Italy.
Belemnitida incertae sedis
(Pl. 2, Figs. 1 a-c, 2 a-c, 3 a-c, 6 a-c, 10 a-c)
Material
Two complete specimens (NS 20/799, /1180), one
specimen lacking the alveolar region (NS 20/772), six
rostra just analysed by Combémorel et al. (1994) coming
from the same levels of the here studied belemnite
assemblage (NS 20/810, /811, /813, /814, /816, /817),
and several rostra lacking the alveolar region (NS
20/769, /842, /858, /859, /861, /920, /917, /918, /900,
/901, /912, /921, /927, /942, /1175, /1185, /1407, /1408,
/1409, /1410, /1411).
Description
Large cylindriconical to slightly hastate, more or less
compressed rostrum. The outline is symmetrical, cylindriconical or weakly hastate; the profile is slightly asymmetrical with a more inflated dorsum. The long flanks
converge rapidly towards the apex. The apical region is
short respect to the total length of rostrum, and ends with
an obtuse and mucronate point. A well marked deep and
broad ventral groove runs from the alveolus to the apex.
The transversal section is more or less compressed.
Some rostra show an epirostrum.
Dimensions
See Tab. 9
Remarks
We assume the same open nomenclature as in
Combémorel et al. (1994a), because the herein analyzed
specimens were collected from the same layers which
gave the fauna studied by the over-mentioned authors.
The specimens show the same features as pointed out by
Combémorel et al. (1994a):
- these rostra exhibit a collection of characters which are
found separately in belemnites belonging to different
genera and families:
- the groove proceeding up to the apex and the
compression of the post-alveolar part of the rostrum are
characteristic of Megateuthididae of the Aalenian and
Bajocian;
- the extremely short termination of the apical region and
the very great width of the groove are characteristic of
the Mesohibolitidae (in particular Pachybelemnopsis),
which first appears in the Lower Bajocian.
For this reason alone it is not possible now to attribute a
genus or family name to these rostra, they need a further
and particular taxonomic-systematic study. Moreover
the individuation of two groups, characterized by rostra
either slender elongate, weakly hastate or more robust
cylindrical, might point out the presence of two taxa.
Stratigraphical range
STRATIGRAPHY
AND PALAEOBIOGEOGRAPHY OF AALENIANEARLY BAJOCIAN BELEMNITE GENERA
In the Lower Jurassic (Hettangian-Pliensbachian) the
order Belemnitida is known from Europe, Turkey,
Greenland, and northern Africa. Then, in the Toarcian,
Belemnitida reached N Siberia producing an endemic
fauna in Toarcian/Aalenian times (Sachs & Nalnjaeva
1970, 1975; Weis & Mariotti, in press). From Siberia
they reached (via the Arctic seas) Arctic Canada, also
forming an endemic fauna (Jeletzky, 1980). Across
Europe, the Early Jurassic belemnite faunas show a
markedly similar composition (Doyle, 1987). The
13
Boreal and Mediterranean provinces
differentiated owing to the Bajocian
transgression which caused the end of
the previous uniformity of the marine
invertebrate faunas.
The genus Holcobelus, especially with
the well-known species H. munieri
and H. blainvillii, appears to be
characteristic of the Aalenian-Early
Bajocian. The overall distribution of
Holcobelus extends from England
(Dorset; Phillips, 1868), to France
(Normandy, Vendée; Eudes-Deslongchamps, 1878; Voltz, 1830; d’Orbigny,
1842-50; Stolley, 1919, 1927; Roger,
1956;), Portugal (Cap Mondego,
Pucanica, Porto de Moz, Ancan;
Choffat, 1880), Luxembourg (Weis &
Mariotti, in press), Germany
(Scheffhen, Gosheim, Plettenberges;
Riegraf, 1980), Austria (St. Veit,
Vienna; v. Hochstetter, 1897),
Switzerland (Bâle; Greppin, 1898), Romania (Lazuri
Valley, Strungarului; Preda, 1975), Bulgaria (Baledie
Han; Stoyanova-Vergilova, 1990), western Turkmenia,
eastern Siberia, Daghestan, Caucasus (Krimholz, 1931,
1958; Noutzubidse, 1966) and southern Italy
(Combémorel et al., 1994a).
Megateuthis and Brevibelus are widespread genera,
particularly abundant in England, France (Normandy),
Luxembourg, Germany, Switzerland, Austria, Romania,
Bulgaria and very rare in southern Italy. The genus
Pachybelemnopsis is recorded from the Lower Bajocian
in England, France, Luxembourg, Germany, Switzerland, Austria, Romania and from the Upper Aalenian in
southern Italy, Bulgaria and Daghestan. Hibolithes is
found in France, Luxembourg, southern Germany and
Poland from the Lower Bajocian on, while in southern
Italy it is recorded in the Upper Aalenian (Weis &
Mariotti, in press).
At the Aalenian-Bajocian boundary a peculiar fauna,
dominated by the genus Holcobelus with several species,
is reported in Normandy (France). A very similar
Holcobelus-dominated fauna is also present in Haute
Provence (France) at Lac du Castillon near Castellane
and at the Truc de Balduc near Mende (Département de
PLATE 1 (page 14)
Fig. 1 - Brevibelus breviformis, NS 20/1184. a) right lateral view, b) ventral view, c) left lateral view; x 1.
Fig. 2 - Holocobelus munieri, NS 20/878. a) right lateral view, b) left lateral view, c) ventral view; x 1.
Fig. 3 - Holcobelus trauthi, NS 20/1181. a) right lateral view, b) left lateral view, c) ventral view; x 1.
Fig. 4 - Holcobelus trauthi, NS 20/1400. a) left lateral view, b) right lateral view, c) ventral view; x 1.
Fig. 5 - Holocobelus munieri, NS 20/838. a) right lateral view, b) left lateral view, c) ventral view; x 1.
Fig. 6 - Holcobelus tetramerus NS 20/773. a) ventral view, b) right lateral view, c) left lateral view; x 1.
Fig. 7 - Holcobelus subblainvillei, NS 20/1404. a) right lateral view, b) left lateral view, c) ventral view; x 1.
Fig. 8 - Holcobelus tetramerus, NS 20/1391. a) right lateral view, b) left lateral view, c) ventral view; x 1.
Fig. 9 - Pachybelemnopsis baculiformis, NS 20 850. a) left lateral view, b) right lateral view, c) ventral view; x 1.
Fig. 10 - Hibolithes wuettembergicus, NS 20/789. a) right lateral view, b) left lateral view, c) ventral view; x 1.
Fig. 11 - Holcobelus tschegemensis, NS 20/847. a) left lateral view, b) right lateral view, c) ventral view; x 1.
Fig. 12 - Megateuthis sp., NS 20/819. a) right lateral view, b) transverse section at the middle part of the alveolar region,
c) transverse section at the apical region; x 2.
Fig.13 - Pachybelemnopsis baculiformis, NS 20/837. a) left lateral view, b) right lateral view, c) ventral view; x 1.
14
PLATE 1
MARIOTTI et al.
PLATE 2
15
16
MARIOTTI et al.
la Lozère) (Riegraf, 1980; Weis & Mariotti, in press).
Therefore the genus Holcobelus appears to mark the
passage from a uniform Europe-wide belemnite fauna in
the Lower Jurassic towards the development of
endemism and the establishment of a Tethyan fauna
(Combémorel et al., 1994a).
In western and central Europe (except Normandy), in the
Lower Bajocian, the predominance of Megateuthididae
(Brevibelus, Megateuthis, Homaloteuthis) and Cylindroteuthididae (Eocylindroteuthis) is noticeable. The
Holcobelidae, with the sole presence of the species
Holcobelus blainvillii, play a minor role compared to
Normandy. In this scenario, the faunas of SW Germany
and Luxembourg show a tight resemblance in the Upper
Aalenian-Lower Bajocian (Weis & Mariotti, in press).
The situation changes in the early Middle Jurassic of the
Mediterranean area. This is characterized by the families
Mesohibolitidae and Holcobelidae and by rare megateuthidids (Megateuthis and Brevibelus) (StoyanovaVergilova, 1990). Pachybelemnopsis is common in
western and central Europe in the Humphriesianum
Zone (Riegraf, 1980; Weis, 2006) being followed by the
earliest Hibolithes. In western and central Europe,
within the Humphriesianum Zone, a faunal changeover
occurs. It is marked by the decline of the suborder
Belemnitina, with Megateuthis and Brevibelus present
until the end of the Bajocian, and the rise of the Tethyan
Pachybelemnopseina with Pachybelemnopsis and
Hibolithes dominating (Weis & Mariotti, in press).
To summarize, in the Late Aalenian belemnite assemblages are characterized:
in Normandy (France) by Holcobelus (very abundant),
Brevibelus (abundant) and the first rare Megateuthis;
in Germany by Holcobelus, Brevibelus and Homaloteuthis;
in Bulgaria by Holcobelus, Brevibelus, Pachybelemnopsis and Paramegateuthis.
These assemblages change in the Early Bajocian:
in Normandy (France) Megateuthis (abundant), Pachybelemnopsis (abundant) and Holcobelus;
in Germany and Luxembourg Megateuthis (abundant),
Brevibelus, Pachybelemnopsis and Eocylindroteuthis;
in Bulgaria Paramegateuthis, Pachybelemnopsis and
Megateuthis.
In southern Italy (Cozzo di Mastro Pasquale, Calabria)
the composition of the belemnite fauna is different in the
Aalenian-Early Bajocian. The following taxa have been
identified: Holcobelus (abundant), Pachybelemnopsis,
Brevibelus (very rare), Megateuthis (very rare), Hibolithes (rare) and Belemnitida incertae sedis (very abundant). This latter group is represented by numerous specimens that may be ascribed to a new taxon whose systematic position apparently lies halfway between
Megateuthididae and Mesohibolitidae; this new systematic group probably is the result of a changing palaeogeography during the early Middle Jurassic. Therefore
the belemnite assemblage from Cozzo di Mastro
Pasquale shows some overall affinities with the faunas
recorded in Normandy (France) and Bulgaria though
with some differences in the relative abundance of the
genera and species.
ACKNOWLEDGEMENTS - We thank Dr. A. Di Cencio for
identification of the ammonites; Prof. R. Matteucci and Prof.
U. Nicosia for the palaeontological discussion and encouragement, Mr. E. Dominici for the photographs and graphics; Mr.
A. Faber (National Museum for Natural History Luxembourg)
for logistical support and Dr. W. Riegraf and Dr. D. Fuchs for
discussions and the loan of comparative material.
This paper is supported by the grant Prin 2006 (E. Turco
scientific leader), and by the research project “Jurassic
coleoids” of the National Museum for Natural History
Luxembourg.
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AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE FROM

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