AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE FROM

INTRODUCTION

The purpose of this paper is to describe in detail apeculiar belemnite fauna, composed of taxa ascribed toHolcobelus Stolley, 1927, Megateuthis Bayle, 1878,Brevibelus Doyle, 1992, Pachybelemnopsis Riegraf,1980, Hibolithes Denis de Montfort, 1808 andBelemnitida incertae sedis. The belemnites were collect-ed from Aalenian-Early Bajocian red marls of theSant’Onofrio formation (Caloveto Group) outcroppingbetween the 8,2 and 8,4 km of the road from Caloveto toBocchiglierio villages (Calabria, southern Italy), in alocality named Cozzo di Mastro Pasquale (Fig. 1). Thesystematic-taxonomic study has allowed some consider-ations about belemnite biostratigraphy and palaeobio-geography. A fauna was first described from this outcropin a preliminary study by Combémorel et al. (1994a).Continued sampling over the last fifteen years producedthe collection described in this paper, representing a fau-nal assemblage that is unique in Italy, but is well knownfrom coeval beds in Romania, Bulgaria, France andsouthern Germany as well as the Caucasus.

PREVIOUS WORK

The European and Mediterranean Aalenian-Bajocianbelemnites are poorly studied either for the lack of out-crops or for the limited number of specimens. The only

AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE FROMPERI-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, [email protected]; [email protected]

°° Robert Weis - Musée National d’Histoire Naturelle de Luxembourg - Section Paléontologie25, rue Münster - L-2160 Luxembourg - [email protected]

Geologica Romana 40 (2007), 1-19

ABSTRACT - A belemnite fauna was collected in southern Italy from hemipelagic marls (Sant’Onofrio forma-tion) 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 wasanalyzed 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-bear-ing levels.

The belemnite fauna is mainly composed of taxa ascribed to the genus Holcobelus (Early Aalenian-MiddleBajocian). This genus is characterized by an apical groove extending from the apex almost reaching the alveolarregion. Although the species Holcobelus trauthi, Holcobelus subblainvillei, Holcobelus munieri, Holcobelustetramerus and Holcobelus tschegemensis are present, the majority of the specimens cannot be identified asbelonging to Holcobelus since their feature set also includes a very short apical region, a long and wide ventralgroove, and a post alveolar compression. These characters are found separately in belemnites belonging to differ-ent 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 Holcobelusand allied forms, the following taxa were also identified: Megateuthis sp., Brevibelus breviformis, Pachy-belemnopsis baculiformis, Hibolithes wuerttembergicus and Hibolithes sp. This fauna has close affinities to fau-nas from Romania, Bulgaria, France, southern Germany, Luxembourg, England and Caucasus.

KEY WORDS: Belemnites, biostratigraphy, Calabria, Aalenian-Early Bajocian.

Fig. 1 - Location map of the examined outcrop (*).

authors who report some of the taxa analyzed here areSchwegler (1938, 1965, 1971) and Riegraf (1980, 1981),who discuss the stratigraphic and geographic distribu-tion of the Jurassic belemnite genera Megateuthis,Mesoteuthis, Eocylindroteuthis, Homaloteuthis, Holco-belus, Brevibelus, Pachybelemnopsis and Hibolithes.Schlegelmilch (1998) figures some original specimensSchwegler (1938, 1965, 1971) that were only knownafter drawings. Althoff (1928) and Waldeck (1975)record Megateuthis from the Bajocian of Bielefeld andLower Saxony (Germany). Kumm (1952) citesHomaloteuthis and Megateuthis from the UpperAalenian 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 belemnitesfrom St. Veit near Vienna (Austria), includingBrevibelus, Megateuthis, Pachybelemnopsis and a newspecies, Belemnites eduardi. Stolley (1927) includes thelatter into his new genus Holcobelus. Phillips (1868) dis-cusses species of Megateuthis, Holcobelus andPachybelemnopsis, but unfortunately no recent, moredetailed studies on these forms are presently available.

From Belgium, Luxembourg and NE France (Alsace-Lorraine) Chapuis & Dewalque (1854) describe for thefirst time Bajocian belemnites (Megateuthis), whileLepsius (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 toHomaloteuthis, Brevibelus, Megateuthis and Pachy-belemnopsis. Weis (2006) describes a Lower Bajocian(Humphriesianum Zone) Megateuthis-Pachybelemno-psis assemblage from Luxembourg. The belemnitemonograph by Eudes-Deslongchamps (1878) is remark-able for NW France (Normandy). The author describesMegateuthis, Brevibelus and Pachybelemnopsis togetherwith several new Upper Aalenian species; unfortunatelyit has not been revised up to now. For these latter formsStolley (1927) institutes the new genus Holcobelus.

Riche (1904), Lissajous (1925) and Roché (1939)record Holcobelus, Brevibelus, Megateuthis andPachybelemnopsis from Central France (Burgundy),while Garnier (1872), Zürcher (1891), Roman &Gennevaux (1912), Gérard (1936) and Riegraf (1980)report Holcobelus and Pachybelemnopsis from theAalenian/Bajocian boundary of Southern France.

In SE Europe the distribution pattern appears to besomewhat more complete. Stoyanova-Vergilova (1982,1985, 1990, 1993) gives an extensive overview ofAalenian and Bajocian belemnites from Bulgaria. Theauthor quotes and figures several species of Holcobelus,Brevibelus, Megateuthis, Paramegateuthis, and Pachy-belemnopsis. A comparable assemblage with Mega-teuthis, Brevibelus, Holcobelus, Pachybelemnopsis andHibolithes is reported by Preda (1975) from the Bajocianof Romania. Such faunas are also reported fromCaucasus and Crimea (Krimholz 1947, 1953). InSouthern Italy, as mentioned above, Combémorel et al.

(1994a) report an isolated belemnite assemblage ofAalenian-Early Bajocian age from Caloveto - the locali-ty which is the object of the present study - withHolcobelus, Brevibelus, Pachybelemnopsis, Hibolithesand Megateuthis.

GEOLOGICAL SETTING

The backbone of the southernmost region of peninsu-lar Italy, Calabria, consists of Early Palaeozoic metamor-phites 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 ofCalabria, named Sila Greca, the Palaeozoic crystallinebasement is transgressively covered by Mesozoic-Palaeogene sediments. This complex was then subjectedto Alpine/Apenninic orogenic deformation, and followedby a new Miocene-Quaternary sedimentary cycle.

The Mesozoic unmetamorphosed sedimentary cover ischaracterized by two distinct successions: the Longo-bucco Group and the Caloveto Group, differing in theirfacies, 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 listof references - to which the reader is referred - docu-menting the early efforts of palaeontologists, mostlyammonite specialists, who tackled the stratigraphy of thearea around the turn of the nineteenth century.Importantly, the papers by Young et al. (1986) andSantantonio & Teale (1987) also introduce a new - as yetinformal - lithostratigraphy, which will be used here.

The Longobucco Group

The Longobucco Group (Triassic-Jurassic boundary toToarcian) (Fig. 2) is exposed across an area of about 150km and it is ~1.5 km thick. It documents sedimentationon a rifted continental margin, in environments evolvingfrom continental (fluvial deposits), to carbonate shelf, toslope and finally to a deep turbiditic basin (see alsoZuffa, 1980).

The Longobucco Group is composed by the followingunits:

1) - Torrente Duno formation (Rhaetian-Hettangian):sandstones (mainly quartzarenites), shales and conglom-erates (red beds) of fluvial environment, 100-200 mthick;

2) - Bocchigliero formation (Sinemurian s.l.): blackishlimestones (wackestones to grainstones), mostly oolitic,with a rich shelly fauna (brachiopods and bivalves), alsowith occasional corals, oncoids and Thalassinoides bur-rows. Material derived from the crystalline basement(e.g. lenses of cross bedded quartz conglomerates) andplant remains (e.g. tree logs) are abundant throughout.Thickness up to about 100 m;

MARIOTTI et al.2 Geologica Romana 40 (2007), 1-19

3) - Fosso Petrone formation (Lower Pliensbachian):bioturbated marls with bivalves, belemnites, ammonitesand plant remains; about 200 m thick; shelf-edge orslope facies;

4) - Fiume Trionto formation (Upper Pliensbachian-Lower Toarcian): mixed siliciclastic/carbonate turbiditesand interbedded hemipelagites, bearing olistoliths, tensto a few hundred metres across, made of the first twounits of the Group. Up to ~1200 m thick.

This succession bears the evidence that the evolutionfrom a continental fluvial environment to a deep marineenvironment must have been driven by rift tectonics inthe Early Jurassic (Santantonio, 1993), ultimately pro-viding the accommodation space for the thick, high sed-imentation-rate Trionto formation turbidites.

Submarine tectonic escarpments exposing the localshelf stratigraphy were in this view the source areas ofthe megaclastic rockfall and debris flow material, andthe turbidites are indeed seen to onlap the Palaeozoicphyllites in several basin-margin localities across theBocchigliero area. The continental deposits (TorrenteDuno formation) gradually evolve to intertidal and tosubtidal marine deposits (Bocchigliero formation).Facies range from quartzose conglomerates and crossbedded hybrid arenites to marls, to cross laminatedoolitic grainstones and bioturbated subtidal wackestonesbearing a diverse marine shelly fauna. The Bocchiglieroformation documents the development of a mixed car-bonate/siliciclastic shelf hosting oolite bars, under theinfluence of river deltas providing abundant siliciclasticinput and plant material sourced by a crystalline hinter-land subjected to a humid climate, and then reworked bytidal and alongshore currents. The transition to the nextunit is documented by subtidal carbonate mud facieswith neritic fauna and flora and the gradual disappear-ance of pebble-and sand-grade siliciclastic material,indicating distal open shelf conditions with decreasingambient energy. Onset of the Petrone formation ismarked by the appearance of cephalopods (belemnitesand ammonites) and the occurrence of thin fine-grainedturbidites, an indication of a deeper-water shelf to slopeenvironment (Young et al., 1986). The lower Toarciantop of the Trionto formation turbidites is cut by modernerosion, so the later Jurassic sedimentary evolution ofthe Longobucco Group is unknown.

The Caloveto Group

The Caloveto Group (Fig. 2) represents a sequencesedimented around and on a basement structural high,and as such spans a long interval of geological time(Early Jurassic to Early Cretaceous) through a muchreduced thickness (maximum 300 m). The LongobuccoGroup acted in the Early Jurassic as a strongly subsidingsediment trap separating the offshore highs of Calovetofrom the Jurassic mainland Calabria. The CalovetoGroup is exposed in small, isolated outcrops dissectedby faults, which often makes bed correlation difficult.

The main outcrops of the Caloveto Group are found intwo main areas: 1. nearby the village of Caloveto, and 2.along the Colognati stream valley. The latter area is thesole having exposures of the younger (Upper Jurassicand Neocomian) part of the succession. The deposits ofthe Caloveto Group display a complex array of uncon-formable contacts and strongly variable geometries andfacies, documenting sedimentation on a rugged sub-strate, under a very strict sinsedimentary tectonic con-trol.

The Caloveto Group is composed by the followingunits, from the base:

1) - Lower Caloveto formation (Pliensbachian). Thiscarbonate unit marks a marine transgression over anintrusive and metamorphic Hercynian basement. It ismade of up to ~80 m of pale-coloured lime grainstones

AALENIAN-EARLY BAJOCIAN BELEMNITE ASSEMBLAGE ... 3Geologica Romana 40 (2007), 1-19

Fig. 2 - Correlated sequences of the Longobucco and Caloveto Groups.Longobucco Group: 1- Torrente Duno Formation; 2- BocchiglieroFormation; 3- Fosso Petrone Formation; 4- Trionto Formation; 5-Hercynian basement (metamorphites); 6- Hercynian basement (grani-toid). Caloveto Group: 1- Lower Caloveto Formation; 2- UpperCaloveto Formation; 3- S.Onofrio Formation (basal red marls); 4-S.Onofrio Formation (pelagic limestone, radiolarites with admixedsiliciclastic and calciclastic turbidites); 5- Hercynian basement (meta-morphites); 6- Hercynian basement (granitoids). Dashed lines are timelines.

and packstones and hybrid arenites, cross-bedded inplaces, bearing rich shallow water benthic faunas (gas-tropods, bivalves, crinoids, corals, Tubiphytes, etc.).

Low-energy cyclic mud-rich facies are remarkablymissing in this unit. This formation also bears an up to40 m thick conglomerate facies, with clast- to matrix(coarse carbonate sand)- supported conglomerates bear-ing rounded pebbles derived from the basement. Theseconglomerates are diachronous, marking the irregular,non conformable contact to the phyllites at Caloveto(more on this below). The change to the next unit can beeither sharp, through a mineralized paraconformity, ortransitional, and made of an up to 10m thick package ofpink brachiopod- (mostly rhynchonellids) to crinoid-richpackstones. The age of the unit is poorly constrained, butthe top levels bear Agerina martana (Farinacci), aPliensbachian benthic foram.

2) - Upper Caloveto formation (Toarcian, Serpentinusto Dispansum Zones), about 0.5-8 m of red pelagic,whole-fossil lime wackstones and nodular marly mud-stone (ammonitico rosso facies). These red pelagitesoften fill deep fractures which cross the former unit aswell as the basement in the form of neptunian dykes.Ammonites, echinoderms (mostly crinoids), bra-chiopods and posidoniid bivalves characterize this unit.A sedimentary hiatus of some 2 My must exist at thePliensbachian-Toarcian boundary, corresponding to theiron-stained basal surface. The ammonite assemblagerecovered indicates the Lower (not lowermost) ToarcianSerpentinus Zone.

3) - Sant’Onofrio formation (Late Toarcian to Haute-rivian), maximum thickness of 150 m, comprising redbioturbated (Zoophychos) (Fig. 3) marls, Aptychus lime-stone, radiolarian cherts and calpionellid limestone.These “background” lithologies host a variety of spec-tacular gravity flow deposits (rock falls, debris flows,turbidites) in the outcrop areas of the high Colognati

stream valley, near Rossano (about 15 km west of theCaloveto outcrop area). These deposits include wide-spread hybrid arenites, clasts of the crystalline basementup to boulder size, and peloid-oolite calcarenites fromunknown carbonate platform source areas. The variouslithologies are, at present, dealt with as informal mem-bers, but may end up being treated as formations, shouldthe Sant’Onofrio be elevated at group rank. Ammonites,belemnites and posidoniid bivalves occur in the redmarls. Apart from aptychi, ammonites are virtuallyabsent in the post-Bajocian sediments, in whichbelemites and other fauna occur sporadically.

The Caloveto Group indicates the initial developmentof carbonate aprons around islands made of Palaeozoicrock. These carbonates have great lateral variability, buttheir ubiquitous grainstone to conglomerate facies is anevidence for high energy conditions in a coastal environ-ment. Coral patches occurred sporadically, but nowheredid an inner lagoon with a low-energy muddy faciesexist. By contrast to the Longobucco basin, that initiallydeveloped as a wide shelf backed by a continent with afluvial system, the Lower Caloveto formation was sedi-mented due to subsidence of emerged land offshore, andthese islands were too small to host a proper drainagesystem. Thence the white (as opposed to black) color ofthe limestone, and the lack of any plant material. Thesecarbonates are locally seen to fill fractures in the phyllitebasement, suggesting that subsidence had a strong tec-tonic component.

Synsedimentary faulting was certainly most active inthe Toarcian, and this probably had two main effects: 1.the foundering and drowning of any shallow water car-bonate system, and 2. the tectonic dissection of both thebasement and the Lower Caloveto limestone. This pro-duced a network of neptunian dykes, and set the condi-tions for the development of various unconformable con-tacts, due to the birth of a markedly irregular submarinetopography. The Colognati area has megaclastic bedssealed by late Toarcian ammonite-rich deposits(Santantonio & Teale, 1987; Santantonio, 1993), sofaulting was active across both the main outcrop areasmentioned above.

The deposits of the Sant’Onofrio formation exhibitvarious types of geometric relationships with their sub-strate. In the Caloveto area contacts vary from paracon-formities to angular unconformities, both with the base-ment and with the faulted shallow-water limestone. At IlTorno (Colognati Valley), Aalenian marls withZoophychos are seen to drape the Toarcian megabrecciaas well as to onlap the granite basement.

The youngest beds seen at Caloveto are the radiolarianchert facies (?Callovian-?Oxfordian), which, where rest-ing unconformably on the Lower Caloveto formation, areseen to induce the growth of chert nodules in the latter,by analogy with basin-margin unconformities in theApennines (Santantonio et al., 1996; Galluzzo &Santantonio, 2002). The Aptychus limestone is found inthe Colognati area, and is seen to change into a calpionel-


Fig. 3 - Zoophychos from red marls (lower part of the Sant’OnorioFormation).

lid limestone (Maiolica facies) which is markedly litho-clastic (basement clasts, plus clasts of the older forma-tions) and also bears turbiditic calcarenites. These fea-tures might be indicative of a reactivation of EarlyJurassic faults around the Tithonian/Berriasian boundary.

The “Cozzo di Mastro Pasquale” outcrop

At km 8.2 along the road connecting the villages ofCaloveto and Bocchigliero, in a locality named “Cozzodi Mastro Pasquale” (Fig. 1), it is possible to observe: a)the top of the Lower Caloveto formation; b) the base ofthe red nodular limestone of the Upper Caloveto forma-tion; c) the marls of the Sant’Onofrio formation.

a) The top of the Lower Caloveto formation is an ero-sional surface, locally iron-stained. One remarkable out-crop at a narrow topographic terrace demonstrates therichly fossiliferous, condensed, nature of the last bed ofthe unit. This bears ostreids, echinoids, gastropods, bra-chiopods, and set in a matrix of hybrid grainstone (crys-talline granules). The first occurrence of rare ammonites(Protogrammoceras sp., Pliensbachian) indicates theonset of a pelagic influence.

b) This area also features the best (though not thethickest) outcrop of the Upper Caloveto formation. Thisis made of red pelagic limestone and marly limestone,with echinoids in the lower part, brachiopods andammonites of Early Toarcian age (Serpentinus Zone).The first centimetres of the Upper Caloveto formationare a red condensed wackestone bearing specimens

ascribed to Hildaites serpentinus. Marly limestone withHildoceras sublevisoni and Mercaticeras sp. (middleToarcian) follows, attaining a maximum thickness of 1m. 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 madeof thick bioturbated beds alternating with thin laminatedposidoniid-rich beds. Soft marl clasts with a thin iron-rich coating could be the product of reworking of incip-ient hardgrounds by burrowers. About 7 m above the topof the Lower Caloveto limestone in the section describedearlier for the Toarcian, which displays about 16 m ofSant’Onofrio marls, a level with large (10-15 cm longeraxis) bivalves probably belonging to the genusInoceramus also produced a rich Aalenian ammonitefauna. 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 foundin situ from 6 m to 11 m above the stratigraphic base ofthe unit. Tmetoceras scissum indicates the earlyAalenian (Opalinum Zone; Venturi 1982; Venturi &Ferri, 2001; Pallini et al., 2004). Erycites intermediusappears at the base of the Opalinum Zone (EarlyAaleniano), while Erycites fallifax is generically anAalenian form. Ancolioceras opalinoides (Mayer) is amarker of the Opalinoides Subzone of the MurchisoneZone. This assemblage therefore covers the middle partof the Opalinum Zone to the middle part of Murchisonae


Fig. 4 - Panoramic view of the Cozzo di Mastro Pasquale outcrop near Caloveto. Dashed line is the trace of the basal unconformity of onlappingmarls (lower part of the Sant’Onofrio Formation).

Zone (Cresta, 2002; ed.). Two stephanoceratids werefound in the upper part of red marls, one in a displacedblock, 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 EarlyBajocian Discites Zone in the Umbria-Marche Apenni-nes. The same species is reported by Callomon &Chandler (1990) from beds of Late Aalenian-EarlyBajocian age in Dorset (Southern England). In exposuresother than those of the Cozzo di Mastro Pasquale area,the lower part of the marl succession has also yieldedammonites indicative of a Late Toarcian (Aalensis Zone)and Early Aalenian (Opalinum Zone) age. The present-day topography, and an unconformable contact withMiocene conglomerates and sandstone, truncate theupper part of the marls, which are seen to change con-formably to red radiolarites a couple of metres above theDocidoceras-bearing bed.

The belemnites studied in this paper were collectedfrom the red marls of the Sant’Onofrio Formation (Fig.6), and their stratigraphic distribution (Aalenian-EarlyBajocian; Combémorel et al., 1994a) is in good agree-ment with ammonite stratigraphy as established in thearea.

An age for the top of the Sant’Onofrio Formation canbe tentatively placed in the Bajocian.

MATERIAL AND METHODS

The terminology herein used is derived from Doyle &Kelly (1988) and Mariotti (2003). Individual descri-ptions also include size measurements in the cases ofcomplete or almost complete specimens. All measuresare in millimeters. Estimated values for incomplete spec-imens are marked by an asterisk (*). Dimensional adjec-tives are used as in Doyle & Kelly (1988); the terms

small, medium and large, related to the length of the ros-trum (L), are respectively referred to L < 80 mm, Lbetween 80 and 110 mm and L > 110 mm.

Abbreviations as in Mariotti (2003): L, total preservedlength; Dv, dorso-ventral diameter at alveolar opening;Dl, lateral diameter at alveolar opening; Dvmax, maxi-mum ventral diameter; Dlmax, maximum lateral diame-ter; X, length from apex to Dmax; Ic, compression index,the ratio between dorsal-ventral diameter and lateraldiameter, calculated at the level of the alveolar opening(Ica), and at the level of the maximum dorso-ventraldiameter (Icm); Id, depression index, the ratio betweenlateral diameter and dorso-ventral diameter calculated atthe level of the alveolar opening (Ida), and at the level ofthe maximum lateral diameter (Idm).

All the collected specimens are stored in thePalaeontological Museum of the Dipartimento diScienze della Terra of the University “La Sapienza” ofRome under collection number NS 20/…

The rostra collected are not in a good state of pre-servation, even if often they are complete. The phragmo-cone is never preserved but frequently the alveolus isobservable. Some of the rostra as well as the ammonites,preserved always as inner moulds, show strong tectonicstress deformation (Fig. 7 a-c).

Furthermore, some specimens exhibit borings made byacrotoracic cirripeds (Fig. 7 d). These organisms aremarine sessile crustaceans with a boring habitat in car-bonate substrata. They form in the substratum a sac-shaped cavity with a comma-shaped opening (with around end and a sharp end), with a total length onlyrarely exceeding 2 mm. These cirripeds orient the cirriagainst the current flux from the rounded portion of theopening directing them towards the sharpened portion(Fig. 8) (Petriconi, 1971). Assuming that borings post-date death of the belemnites and settlement of the ros-trum on the sea-bottom, it is conceivable that sedimenta-


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; dTmetoceras scissum, Aalenian, Opalinum Zone, x 1.

tion rates were as slow as to permit the colonisation ofrostra before they were covered by sediment. Sometimesthe borings reach the innermost part of the rostrum (Fig.7 d).

SYSTEMATICS

The classification adopted here is based on Riegraf(1995) and Riegraf et al. (1998).

Order Belemnitida MacGillivray, 1840Suborder Belemnitina MacGillivray, 1840Family Megateuthididae Sachs & Nalnjaeva, 1967

Genus Brevibelus Doyle, 1992

Diagnosis“Small, short and robust, cylindriconical to conicalMegateuthididae. Outline symmetrical and conical tocylindriconical, profile symmetrical to slightly asym-metrical, otherwise similar to the outline. Apex obtuse tomoderately acute, often mucronate. Venter inflated insome species. Transverse sections quadrate, compressedin some species, depressed in others. Apex devoid ofgrooves or striae. Lateral lines may be well-developed,consisting of two weak and parallel depressions separat-ed by a relatively well-developed ridge. The phragmo-cone is ventrally displaced, penetrating one half to onethird of the rostrum. Apical lines strongly cyrtolineate,apical angle approximately 27°.” (After Doyle, 1992)

Type speciesBelemnites breviformis Voltz, 1830

RemarksParapassaloteuthis Riegraf, 1980 is the only genus sim-ilar to Brevibelus, but the latter differs in having a moreconical rostrum, no apical groove and weaker laterallines.

Brevibelus breviformis (Voltz, 1830)

(Pl. 1, Fig.1 a-c)


Fig. 6 - Belemnites bearing red marls of the lower part of theSant’Onofrio Formation. The white arrow indicates a hammer.

Fig. 7 - a-c) boudinaged rostra, x 1; d) borings of acrotoracic cirripedesat the outer surface (left) and in the innermost part of the guard, x 1.

Fig. 8 - The boring of an acrotoracic cirriped (after Petriconi, 1971).

*1830 Belemnites breviformis Voltz, p. 42, Pl. II, Figs. 2-4.1992 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, 27-29

MaterialTwo complete juvenile specimens in mediocre state ofpreservation (NS 20/877-/1184).

DescriptionThe small rostrum is typically conical and slender. Theoutline is symmetrical and conical, the profile is nearlysymmetrical with a slightly flattened venter. The apex ismoderately acute. The transverse section is subquadratein the alveolar region and becomes elliptical in the api-cal region. No groove is present. The phragmocone pen-etrates one half of the rostrum.

DimensionsSee Tab. 1

RemarksThe characteristic conical and slender shape and the lackof groove permit a systematic attribution to a juvenileform of Brevibelus. The juvenile stadium of the twospecimens is based on ontogenetic observations of Weis(1999, p. 224). The only similar species is Brevibelusgingensis, but Brevibelus breviformis is different for itsmore slender shape. In Italy this species, common in the European belemniteassemblages, is recorded only from the “Cozzo diMastro Pasquale” outcrop.

Geographical and stratigraphical distributionUpper Toarcian to Lower Bajocian of Europe, Russia,western Canada (Doyle, 1992), and Aalenian-LowerBajocian of southern Italy.

Genus Megateuthis Bayle, 1878

DiagnosisVery large conical to elongate cylindriconical Megateu-thididae, with a well-developed epirostrum. The outlineand profile are symmetrical and conical to cylindriconi-cal. The apex is obtuse and bears dorso-lateral groovesand generally well-marked striae. Transverse sectionsare compressed and elliptical.

Type speciesBelemnites giganteus v. Schlotheim, 1820 (= BelemnitesAalensis Voltz, 1830) by subsequent designation(Douvillé, 1879, p. 91)

Megateuthis sp.

(Pl. 1; Fig. 12 a-c)

MaterialOne juvenile fragment showing the alveolar and partiallythe stem region (NS 20/819).

DescriptionThe small conical rostrum is characterized by two apicaldorso-lateral grooves visible on the transverse section. Itlacks a ventral groove and is strongly compressed withan elliptical cross section. The alveolus probably pene-trates one fourth of the rostrum.

RemarksThe fragmentary preservation does not permit a specificattribution. The blunted apical part of the rostrum expos-es the typical juvenile stages of Megateuthis with clearindications of two apical dorso-lateral grooves. BothSchwegler (1965) and Pugacewska (1961) illustrate sim-ilarly short, compressed and conical juveniles ofMegateuthis elliptica from the Bajocian of Poland andsouthern Germany. The fragmentary state (lack of apicalregion) of the unique collected specimen does not allowdrawing any further comparison.

Geographical and stratigraphical distributionBajocian of Europe, ?North America (Doyle, 1992), andAalenian-Lower Bajocian of southern Italy.

Suborder Pachybelemnopseina Riegraf, 1998Family Holcobelidae Gustomesov, 1977

Genus Holcobelus Stolley, 1927

DiagnosisSmall to medium sized, elongated, conical to cylindricalHolcobelidae. Apex acute or obtuse (orthorostrum). Noapical grooves. Intermediate ventral groove extendingfrom the alveolar region to nearly the apex. Transversesections rounded or elliptical, compressed. Epirostrumwell-developed in some species.

RemarksThe phylogenetic affinities of Holcobelus werediscussed in Combémorel et al. (1994a).

Type speciesBelemnites munieri Eudes-Deslongchamps, 1878.

Holcobelus trauthi Stolley, 1927

(Pl. 1, fig. 3 a-c)


*1927 Holcobelus trauthi Stolley, p.118, pl. XXIV, fig. 7-8. 1994 Holcobelus trauthi - Combémorelet al., p. 47, Pl. 1, fig. 4.

MaterialSix complete rostra (NS 20/1181,/1179, /862, /1178, /1400, /1403), onejuvenile form (NS 20/828), one speci-men lacking the alveolar region (NS20/815), and several fragments of thedistal part of the alveolar region (NS20/786, /823, /776, /866).

DescriptionThe middle sized rostrum is conical inthe posterior. Outline and profile aresymmetrical and cylindriconical, veryslightly hastate in the adult. The ventral groove is narrowand well-marked, reducing towards the apex which itappears to meet. The transverse section is sub-rectangu-lar and compressed. The apical region is slightly obtuse.


RemarksThis species is similar to the original specimens ofStolley (1927, pl. XXIV, fig. 7-8). The closest species isHolcobelus subblainvillei but the latter shows a moreconical slender shape and less compression.

Geographical and stratigraphical distributionLower Bajocian of Normandy (France), and Aalenian-Lower 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.

MaterialFive complete rostra (NS 20/897, /1404, /785, /784,/880) and several fragments.

DescriptionThe conical rostrum is medium-sized and slender. Theprofile is symmetrical and conical; the outline issymmetrical, but less conical than profile. A ventralnarrow deep groove extends from the alveolus next tothe apex. The apical region ends with an acute andslightly mucronate point. The transverse section issubcircular to slightly compressed. The phragmoconepenetrates one third of the rostrum.


RemarksThis species is very similar to Holcobelus blainvillii, butit differs by its groove extending from alveolus to apex.Another taxon comparable with Holcobelus subblain-villei is Holcobelus munieri. Holcobelus munieri has avery long and more slender rostrum and an apical regionmore acute.Combémorel et al. (1994b), in the revision ofd’Orbigny’s collection, figured a longitudinal section ofHolcobelus munieri with an epirostrum, and retainedthat Holcobelus subblainvillei would be Holcobelusmunieri without epirostrum. This might be truth, but forthe moment we do not have appropriate material for fur-ther investigation.

Geographical and stratigraphical distributionUpper Aalenian and Lower Bajocian of Normandy,Haute-Provence (France), and Aalenian-Lower Bajocianof 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.)


MaterialTwo juveniles (NS 20/880, /843), two specimens lackingthe distal end (NS 20/838, /768) and one specimenlacking the most part of the alveolar region (NS 20/878).

DescriptionVery slender conical and slightly compressed rostrum.The outline and profile are symmetrical and conical. Thedorsal, the ventral sides and the flanks converge weaklybut continuously till the apex. The apical region is acuteand the end is sometimes very slightly obtuse. A deepventral groove starts from the alveolar opening, persistsinto the stem region disappearing at the apex. The trans-versal section is weakly compressed, subquadrate. Thephragmocone penetrates one fourth of the rostrum. Theadult stage presents an epirostrum.

DimensionsSee Tab. 4Remarks

This species differs from the other species ascribed toHolcobelus by the long slender conical rostrum, theconical apical region and the long groove.

Geographical and stratigraphical distributionUpper Aalenian and Lower Bajocian of Normandy,southern France, Bulgaria, and Aalenian-Lower Bajo-cian 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.

MaterialFive complete specimens (NS 20/774, /773, /892, /1399,/1405).

DescriptionThe rostrum is small-sized and compressed. The outlineis symmetrical and conical, the profile is symmetricaland cylindriconical. The flanks and the ventral and later-al sides run parallel to converge suddenly in the apicalregion ending with an obtuse point; this is evident in the

adult specimens. The transversal section is compressedand subrectangular. A deep ventral groove extends fromthe alveolus till the apex. The phragmocone, slightlyeccentric ventrally, penetrates for one third of the totallength of the rostrum.


RemarksThe small-sized rostrum, the compressed subrectangulartransversal section and the obtuse apical region makethis species unique within the genus Holcobelus.

Geographical and stratigraphical distributionUpper Aalenian of Normandy, Haute-Provence (France)and Aalenian-Lower Bajocian of southern Italy.

Holcobelus tschegemensis (Krimholz, 1931)

(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.

MaterialTwo juvenile specimens (NS 20/824, NS 20/851), onespecimen lacking the distal end (NS 20/775), onespecimen lacking the apical region (NS 20/1406), threefragments of the apical region (NS 20/864, /907, /1412),one rostrum lacking the terminal end and part of thealveolar region (NS 20/865) and two specimens withoutthe apical region (NS 20/847, /777).

DescriptionThe medium-sized rostrum is typically slender and thin.The profile and outline are symmetrical and cylindrical.The transverse section is elliptical to subquadrate andweakly compressed for all the length of rostrum. Thealveolus penetrates one sixth of the rostrum. The longventral groove is deep in the anterior region, and fadesslightly out towards the apex.



RemarksThis species, characterized by a medium-sized slenderan thin rostrum and a long ventral groove, is outstandingin the genus Holcobelus.

Geographical and stratigraphical distributionUpper Aalenian of the Caucasus, Bulgaria, HauteProvence (France) and Aalenian-Lower Bajocian ofsouthern Italy.

Family Mesohibolitidae Nerodenko, 1983

Genus Hibolithes Denys de Montfort, 1808

DiagnosisElongated, slender Mesohibolitidae. Both profile andoutline symmetrical and usually hastate. Maximumtransverse diameter usually at the apical region.Transverse section generally circular, in some casesdepressed in the apical region in some others com-pressed in the alveolar region. Shallow ventral canal lim-ited at the anterior half of the rostrum. Well evident lat-eral lines. Central apical line.

Type speciesHibolithes hastatus Denys de Montfort, 1808

Hibolithes wuerttembergicus (Oppel, 1856)

(Pl. 1, Fig. 10 a-c; Pl. 2, Figs. 4 a-c, 8 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. 67-70, 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.

MaterialThree juvenile specimens (NS 20/788, /789, /1401) allwithout phragmocone and alveolus.

DescriptionSmall-sized fusiform rostrum with outline and profilesymmetrical and fusiform. The maximum width of ros-trum occurs between 1/3 and 1/4 from the apex. A verythin and narrow ventral canal is present in the most prox-imal part. Weak lateral lines are visible on the flanks.The transverse section is more or less circular.


RemarksThe two juvenile specimens are ascribed to Hibolitheswuerttembergicus by their fusiform shape, the weakalveolar canal and lateral lines. They are synonymouswith the juveniles figured by v. Zieten (1830-33, Pl. 25,fig. 3c-f). Pugaczewska (1961) describes and figuresthree ontogenetic stages for this species: nepionic, nean-ic and ephebic-gerontic. The here analyzed specimenscould be nepionic forms for their fusiform shape, thelength and the tapering of the terminal portion and thepoorly incised alveolar canal.

Stratigraphical range and geographical distributionBajocian to Bathonian of Germany, Luxembourg,Poland and Aalenian-Lower Bajocian of southern Italy.

Hibolithes sp.

(Pl. 2, Fig. 5 a-c)

MaterialThree stem fragments close to the alveolar region (NS20/779, /1395, 848) and one fragment showing thealveolar region and partially the stem region (/1413).

DescriptionFragments of medium and large rostra with a probablefusiform shape. A clear broad ventral canal is observ-able. The dorsal side shows a short depression extendingdistally as a flattened area. The transverse sections arecircular to slightly elliptical. Two lateral lines are presenton each flank.

RemarksThe type of the canal, the transverse section, the pres-ence of lateral lines and a short dorsal depression permit


to ascribe these fragments to Hibolithes.

Stratigraphical range and geographical distributionAalenian-Lower Bajocian of southern Italy.

Genus Pachybelemnopsis Riegraf, 1980

DiagnosisCylindrical to hastate, elongate Mesohibolithidae.Outline symmetrical. Profile asymmetrical. Apex acute.Transverse section generally depressed in the apical andstem regions. Broad, ventral alveolar canal sometimesextending up to the apex. Apical line ventrally eccentric.Lateral lines rarely present.

Type speciesBelemnites 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) baculiformisRiegraf, p. 179-181, text-figs. 160-161, Pl. 1, fig. 11[cum syn.]1998 Belemnopsis baculiformis - Schlegelmilch, p. 79,Pl. 16, figs. 10-11v. 2006 Pachybelemnopsis baculiformis - Weis, p. 161,fig. 12

MaterialOne juvenile specimen (NS 20/830), three specimenslacking the alveolar region (NS 20/837, /850, /834), andtwo fragments of the stem region (NS 20/839, /822).

DescriptionMedium sized rostrum with a fusiform shape narrowingdistally to end by a mucronate tip. The transverse sectionis subcircular in the alveolar region and more or lessdepressed towards the apex. A deep ventral canalextends towards the point, it is broad with clear ridgesand its thickness decreases distally; it fades out at thebeginning of the apical region. The lateral lines areweak. The outline is hastate, with maximum diameterlocated at the 2/3 of the total preserved length. Theprofile is hastate, slightly asymmetrical. The apicalregion is short and slightly obtuse.


RemarksPachybelemnopsis baculiformis distinguishes from otherspecies of Pachybelemnopsis by its little club-like formand the relatively long and broad ventral canal.

Stratigraphical range and geographical distributionBajocian (Humpresianum Zone) south-western Germany,north-eastern France, Luxembourg and Aalenian-LowerBajocian of southern Italy.

Belemnitida incertae sedis

(Pl. 2, Figs. 1 a-c, 2 a-c, 3 a-c, 6 a-c, 10 a-c)

MaterialTwo complete specimens (NS 20/799, /1180), onespecimen lacking the alveolar region (NS 20/772), sixrostra just analysed by Combémorel et al. (1994) comingfrom the same levels of the here studied belemniteassemblage (NS 20/810, /811, /813, /814, /816, /817),and several rostra lacking the alveolar region (NS20/769, /842, /858, /859, /861, /920, /917, /918, /900,/901, /912, /921, /927, /942, /1175, /1185, /1407, /1408,/1409, /1410, /1411).

DescriptionLarge cylindriconical to slightly hastate, more or lesscompressed rostrum. The outline is symmetrical, cylin-driconical or weakly hastate; the profile is slightly asym-metrical with a more inflated dorsum. The long flanksconverge rapidly towards the apex. The apical region isshort respect to the total length of rostrum, and ends withan obtuse and mucronate point. A well marked deep andbroad ventral groove runs from the alveolus to the apex.The transversal section is more or less compressed.Some rostra show an epirostrum.


RemarksWe assume the same open nomenclature as inCombémorel et al. (1994a), because the herein analyzedspecimens were collected from the same layers whichgave the fauna studied by the over-mentioned authors.The specimens show the same features as pointed out byCombémorel et al. (1994a):- these rostra exhibit a collection of characters which arefound separately in belemnites belonging to differentgenera and families:- the groove proceeding up to the apex and thecompression of the post-alveolar part of the rostrum arecharacteristic of Megateuthididae of the Aalenian andBajocian;- the extremely short termination of the apical region andthe very great width of the groove are characteristic ofthe 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 furtherand particular taxonomic-systematic study. Moreoverthe individuation of two groups, characterized by rostraeither slender elongate, weakly hastate or more robustcylindrical, might point out the presence of two taxa.

Stratigraphical rangeAalenian-Lower Bajocian of southern Italy.

STRATIGRAPHYAND PALAEOBIOGEOGRAPHY OF AALENIAN-

EARLY BAJOCIAN BELEMNITE GENERA

In the Lower Jurassic (Hettangian-Pliensbachian) theorder Belemnitida is known from Europe, Turkey,Greenland, and northern Africa. Then, in the Toarcian,Belemnitida reached N Siberia producing an endemicfauna in Toarcian/Aalenian times (Sachs & Nalnjaeva1970, 1975; Weis & Mariotti, in press). From Siberiathey reached (via the Arctic seas) Arctic Canada, alsoforming an endemic fauna (Jeletzky, 1980). AcrossEurope, the Early Jurassic belemnite faunas show amarkedly similar composition (Doyle, 1987). The

Boreal and Mediterranean provincesdifferentiated owing to the Bajociantransgression which caused the end ofthe previous uniformity of the marineinvertebrate faunas. The genus Holcobelus, especially withthe well-known species H. munieriand H. blainvillii, appears to becharacteristic of the Aalenian-EarlyBajocian. The overall distribution ofHolcobelus extends from England(Dorset; Phillips, 1868), to France(Normandy, Vendée; Eudes-Deslong-champs, 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 (LazuriValley, Strungarului; Preda, 1975), Bulgaria (BaledieHan; 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 genusPachybelemnopsis is recorded from the Lower Bajocianin England, France, Luxembourg, Germany, Switzer-land, Austria, Romania and from the Upper Aalenian insouthern Italy, Bulgaria and Daghestan. Hibolithes isfound in France, Luxembourg, southern Germany andPoland from the Lower Bajocian on, while in southernItaly 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 similarHolcobelus-dominated fauna is also present in HauteProvence (France) at Lac du Castillon near Castellaneand 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.


PLATE 1


PLATE 2

la Lozère) (Riegraf, 1980; Weis & Mariotti, in press).Therefore the genus Holcobelus appears to mark thepassage from a uniform Europe-wide belemnite fauna inthe Lower Jurassic towards the development ofendemism and the establishment of a Tethyan fauna(Combémorel et al., 1994a).In western and central Europe (except Normandy), in theLower Bajocian, the predominance of Megateuthididae(Brevibelus, Megateuthis, Homaloteuthis) and Cylindro-teuthididae (Eocylindroteuthis) is noticeable. TheHolcobelidae, with the sole presence of the speciesHolcobelus blainvillii, play a minor role compared toNormandy. In this scenario, the faunas of SW Germanyand Luxembourg show a tight resemblance in the UpperAalenian-Lower Bajocian (Weis & Mariotti, in press). The situation changes in the early Middle Jurassic of theMediterranean area. This is characterized by the familiesMesohibolitidae and Holcobelidae and by rare mega-teuthidids (Megateuthis and Brevibelus) (Stoyanova-Vergilova, 1990). Pachybelemnopsis is common inwestern and central Europe in the HumphriesianumZone (Riegraf, 1980; Weis, 2006) being followed by theearliest Hibolithes. In western and central Europe,within the Humphriesianum Zone, a faunal changeoveroccurs. It is marked by the decline of the suborderBelemnitina, with Megateuthis and Brevibelus presentuntil the end of the Bajocian, and the rise of the TethyanPachybelemnopseina with Pachybelemnopsis andHibolithes dominating (Weis & Mariotti, in press).To summarize, in the Late Aalenian belemnite assem-blages are characterized:in Normandy (France) by Holcobelus (very abundant),Brevibelus (abundant) and the first rare Megateuthis;in Germany by Holcobelus, Brevibelus and Homalo-teuthis;in Bulgaria by Holcobelus, Brevibelus, Pachybelemno-psis and Paramegateuthis.

These assemblages change in the Early Bajocian:in Normandy (France) Megateuthis (abundant), Pachy-belemnopsis (abundant) and Holcobelus;in Germany and Luxembourg Megateuthis (abundant),Brevibelus, Pachybelemnopsis and Eocylindroteuthis;in Bulgaria Paramegateuthis, Pachybelemnopsis andMegateuthis.In southern Italy (Cozzo di Mastro Pasquale, Calabria)the composition of the belemnite fauna is different in theAalenian-Early Bajocian. The following taxa have beenidentified: Holcobelus (abundant), Pachybelemnopsis,Brevibelus (very rare), Megateuthis (very rare), Hibo-lithes (rare) and Belemnitida incertae sedis (very abun-dant). This latter group is represented by numerous spec-imens that may be ascribed to a new taxon whose sys-tematic position apparently lies halfway betweenMegateuthididae and Mesohibolitidae; this new system-atic group probably is the result of a changing palaeo-geography during the early Middle Jurassic. Thereforethe belemnite assemblage from Cozzo di MastroPasquale shows some overall affinities with the faunasrecorded in Normandy (France) and Bulgaria thoughwith some differences in the relative abundance of thegenera and species.

ACKNOWLEDGEMENTS - We thank Dr. A. Di Cencio foridentification of the ammonites; Prof. R. Matteucci and Prof.U. Nicosia for the palaeontological discussion and encourage-ment, 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 fordiscussions and the loan of comparative material.

This paper is supported by the grant Prin 2006 (E. Turcoscientific leader), and by the research project “Jurassiccoleoids” of the National Museum for Natural HistoryLuxembourg.


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