PINARD J.D., WEIS R., NEIGE P., MARIOTTI N. & DI CENCIO A. (2014) - Belemnites from the Upper...

Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous (Causses France)

Jean-Daniel Pinard Robert Weis Pascal Neige Nino Mariotti and Andrea Di Cencio

With 9 figures

Abstract Jurassic belemnites represent promising proxies especially for palaeoecological and also palaeobiogeographical reconstructions However basic knowledge on taxonomic composition bios-tratigraphy and diversity of Early Jurassic belemnites is still dramatically low especially for France The present study provides new data on belemnites from the southern part of the Northwest European Province More than 700 specimens have been collected along the Tournadous section in the Causses Basin (Southern France) they represent ten genera and 30 species belonging to Hastitidae Mega-teuthididae Passaloteuthididae and Salpingoteuthididae Furthermore numerous ammonites have been collected allowing to establish a standard chronostratigraphical scheme of the studied section and thus to assign for the first time a precise chronostratigraphical distribution to the Early Jurassic belemnites of Southern France The analysis of belemnite diversity and its variations along the sec-tion compared to the well-known fauna of the northern part of the Northwest European province (Germany and England) open interesting prospective for belemnites palaeobiogeography

Key words Belemnite systematics palaeobiogeography Pliensbachian Toarcian Southern France

1 Introduction

Belemnites ss originated at the beginning of the Early Jurassic (Doyle 1994 RiegRaf 1996a Weis amp Del-sate 2006) though recent findings from Japan pro-vide arguments for a Triassic origin (iba et al 2012) They display a successful diversification during the Jurassic with a peak in species richness for the Toar-cian Detailed monographs have recently been pub-lished providing taxonomic and stratigraphic revised data (eg Doyle 1990 1992 RiegRaf et al 1984 sanDeRs et al 2013 Weis amp MaRiotti 2007 Weis et al 2012) However these modern taxonomic revisions are based on studies that are far from being homogene-ous considering their palaeogeography The Toarcian (Early Jurassic) which concentrates most modern be-lemnite taxonomic revisions is exemplary It is an im-portant period of taxonomic modification for belem-nites Doylersquos synthesis paper (1994) points out that

the Toarcian is also an important period of change in belemnite distribution and concludes that a broad cor-relation between European areas and a notable ende-mism for Siberia occurs However his study was at the generic level and focuses on the northern part of the Northwest European Province and on Siberia Finally and even today modern data for Toarcian belemnites are largely concentrated on the northern part of the Northwest European Province (Germany Great Brit-ain Luxembourg etc) whereas little is known from the southern part of the Northwest European Pro- vince (Causses and Quercy basins Provence Platform northern Spain etc) and from the Mediterranean Province (Italy Austria Southern Spain North Africa etc) In all these areas data are still dramatically low and what is known about belemnites is whether old and without any figuration or sparse (eg some isolat-ed specimens figured see lissajous 1925 sciau 1993 2011 coMbeacuteMoRel in Rulleau et al 1998) This lack

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DOI 1011270077-774920140421 0077-774920140421 $ 575

N Jb Geol Palaumlont Abh 2732 (2014) 155ndash177 ArticleStuttgart August 2014

Kolumnentitel links Jean-Daniel Pinard et al

Kolumnentitel rechts Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous

N Jb Geol Palaumlont Abh ArticleStuttgart August 2014

Achtung Bilder 6-8 unbedingt Originalgroumlszlige

E E

156 Jean-Daniel Pinard et al

of data is due to two factors (1) the low number of belemnites study in these areas and (2) some outcrops (ie in Italy and Spain) are hardened carbonates lead-ing a very difficult extraction of well-preserved speci-mens It is therefore impossible at the present day to portray the Jurassic belemnite diversification at the species level mostly because we lack data from some palaeogeographic areas unless introducing strong pal-aeogeographic biases

In the present study we report a belemnite fauna of more than 700 specimens from the Tournadous section in the Causses Basin This fauna constitutes then a homogeneous and comprehensive record of the Late Pliensbachian and Toarcian belemnite fauna in the southern part of the Northwest European Prov-ince These specimens have been collected in a com-mon field study associating ammonites and belemnites scholars It allows to assign the stratigraphic distribu-

tion of belemnites to a standard chronostratigraphic frame based on ammonites The reported belemnites substantially increase the knowledge of Early Jurassic faunas from the southern part of the Northwest Euro-pean Province a step that could later lead to further palaeogeographic explorations of the Jurassic radia-tion

2 Geological and palaeogeographic settings

We study here a single section (Tournadous) which ex-poses Lower Jurassic marls located in Central-South France (Fig 1) This section has been studied by sev-eral authors principally for its ammonoids content (eg MonestieR 1921 1931 guex 1972 MeisteR 1989 MoRaRD 2004) or to explore environmental changes at the Upper Pliensbachian-Lower Toarcian bound-

Fig 1 Palaeogeographic and geographic setting of Tournadous (Causses France) The provinces boundary is determined on ammonite fauna (Palaeogeographic map modified from DeRa et al 2010)

Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 157

ary (Mailliot et al 2009) The studied portion in the present analysis covers the uppermost Pliensbachian and continues up to the top of the Toarcian (see below) This section belongs to the Causses Basin consid-ered as a small intracratonic basin that was part of an epicontinental sea The basin was stuck between the ldquoMassif Centralrdquo lands on the North and the ldquoMon-tagne Noirerdquo lands on the South Detailed analysis of the structure of the basin sequence stratigraphic con-text and stratigraphical correlations or geochemical

proxies may be found in an extensive literature (eg tRuumlMpy 1983 gRaciansky et al 1998 Mailliot et al 2009 Van De schootbRugge et al 2013) Subsidence is more or less pronounced within the basin leading to variable thickness of the different formations and even to hiatuses (see tRuumlMpy 1983 gRaciansky et al 1998) The studied section (Tournadous) is considered as very proximal (Mailliot et al 2009)

The studied interval covers three successive local formations At the base of the section the ldquoMarnes de

Fig 2 Stratigraphic repartition of collected ammonites and chronostratigraphic scheme of the section (Part 1)




Fig 4 Stratigraphic repartition of the collected belemnites


Villeneuverdquo formation displays dark marls with a typi-cal three nodular beds succession at its topmost part The last nodular bed has been considered to mark the top of the Pliensbachian (Mailliot et al 2009) Fossils

within this formation are generally well-preserved Belemnites frequently bear their phragmocone and ammonites are pyritized Above is the well-known ldquoSchistes cartonsrdquo formation It consists in finely lami-

Fig 5 Belemnites from Tournadous Hastitidae The specimens are all in natural size except where stated otherwise A ndash Pleurobelus compressus (stahl 1824) BEL-256 rostrum in ventral view (A1) lateral view (A2) and dorsal view (A3) B ndash Pleurobelus compressus (stahl 1824) BEL-256 rostrum in ventral view (B1) lateral view (B2) and dorsal view (B3) C ndash Pleurobelus aff compressus (stahl 1824) BEL-258 rostrum in ventral view (C1) lateral view (C2) and dorsal view (C3) D Pleurobelus aff compressus (stahl 1824) BEL-259 medial section of the rostrum (enlargement 2x) E ndash Pleurobelus lagenaeformis (haRtMann in V Zieten 1832) BEL-260 rostrum in ventral view (E1) lateral view (E2) and dorsal view (E3) F ndash Pleurobelus lagenaeformis (haRtMann in V Zieten 1832) BEL-272 medial section of the rostrum (enlargement 2x) G ndash Pleurobelus lagenaeformis (haRtMann in V Zieten 1832) BEL-260 rostrum in ventral view (G1) lateral view (G2) and dorsal view (G3) H ndash Pleurobelus subirregularis (lissajous 1927) BEL-260 rostrum in ventral view (H1) lateral view (H2) and dorsal view (H3) I ndash Pleurobelus subirregularis (lissajous 1927) BEL-260 rostrum in ventral view (I1) lateral view (I2) and dorsal view (I3) J ndash Rhabdobelus donovani RiegRaf in RiegRaf et al 1998 BEL-299 rostrum in ven-tral view (J1) lateral view (J2) and dorsal view (J3) K ndash Rhabdobelus donovani RiegRaf in RiegRaf et al 1998 BEL-300 rostrum in ventral view (K1) lateral view (K2) and dorsal view (K3) L ndash Rhabdobelus aff donovani RiegRaf in RiegRaf et al 1998 BEL-293 rostrum in ventral view (L1) lateral view (L2) and dorsal view (L3)


nated shale sometimes containing silty material Most ammonites are compressed along laminations Rare pyritised nuclei may be found Finally the ldquoMarnes de Fontaneillesrdquo formation expands up to the Aalenian It consists mainly of marls with rare intercalated car-bonate beds Within the Tournadous section a very distinctive calcareous bed (bed number 100 here see Fig 2) occurs at nearly 20 meters above the base Am-monites from this formation are pyritised at the base and calcareous and sometimes slightly compressed at the top Within this ldquoMarnes de Fonteneillesrdquo forma-tion gastropods and rare bivalves have been found Belemnites are frequent well-preserved mostly with their phragmocone

Based on ammonites palaeogeographic distribu-tions elMi et al (1997) considered the Causses Basin to be southern part of the Northwest European Prov-ince whereas for page (2003) it was part of a Submedi-terranean Province for the beginning of the Toarcian (Serpentinum and Bifrons chronozones) and part of the Northwest European Province after (from Variabi-lis to Aalensis chronozones)

3 Chronostratigraphy

Ammonites have been precisely located along the sec-tion (Figs 2-3) From these distributions a chronos-tratigraphic framework is proposed at the chronozone and subchronozone scales (Figs 2-4) For that we ex-plicitly refer to the landmark publications of DoMMeR-gues et al (1997) elMi et al (1997) and page (2003) Chronostratigraphic zonation expressed here (Figs 2-4) thus corresponds to a standard zonation scheme where chronozones (and subchronozones) are defined only at their base (see page 2003) Question marks in the present chronostratigraphic framework denote our impossibility to correctly date the referring beds due to the absence of ammonites The choice of a stand-ard (ie chronostratigraphic) zonation is opportune because when established this framework will serve as a reference to document temporal distribution of belemnite species

According to that approach the studied part of Tournadous section expands from the Margaritatus Chronozone (Upper Pliensbachian) to the Aalensis Chronozone (Upper Toarcian) We were not able to observe the Spinatum Chronozone (last Pliensbachian zone) (Fig 2) However MeisteR (1989) for different sections in the Causses clearly recognizes it Therefore we probably missed index species or associated corre-lating fauna of the uppermost part of the Pliensbachian

which is thus probably present in the Tournadous sec-tion All Toarcian chronozones are documented along the section except for the Tenuicostatum Chronozone (lowermost chronozone of Toarcian stage) This fits well previously published papers which claim that the entire Causses Basin lacks deposits from this lower-most Toarcian interval (tRuumlMpy 1983 guex et al 2001 Mailliot et al 2009) Within the section we note a distinctive condensed interval (bed 100 Figs 3-4) with the existence of fossil accumulation dated from the Thouarsense Chronozone and from the base of the Dispansum Chronozone (Insigne Subchrono-zone) Interestingly this accumulation at that age is recorded by gRaciansky et al (1998 617) in the much closed Quercy Basin but not ndash as far as we know ndash in other Causses Basin sections This is probably due to the relatively proximal position of the Tournadous sec-tion contrary to the other more distal sections of the basin

4 Systematic palaeontologyThe systematics herein used is derived from Doyle (1994) and RiegRaf (2000) Terminology follows Doyle amp kelly (1988) All studied belemnites are housed at the National Museum of Natural History Luxembourg (MNHNL) Col-lected ammonites are housed at the University of Burgundy France (UBGD coll neige and pinaRD) The belemnite ma-terial is composed of 710 specimens including 131 indeter-minate rostra 115 determinate only at the genus level and 464 at the species level Detailed stratigraphic repartition of the different species is given on Fig 4

Subclass Coleoidea batheR 1888Order Belemnitida Zittel 1895

Suborder Belemnitina Zittel 1895Family Hastitidae naef 1922Genus Pleurobelusnaef 1922

Type species Belemnites compressus stahl 1824 Upper Pliensbachian Amaltheenton Formation South-west Ger-many

Pleurobelus compressus (stahl 1824)Fig 5A B

1824 Belemnites compressus sp n ndash stahl p 331994 Pleurobelus compressus (stahl 1824) ndash fischeR p

13 pl 2 fig 31998 Pleurobelus compressus (stahl 1824) ndash schlegel-

Milch p 54 pl 4 fig 3


Material Thirty-six rostra from level 85 Upper Pliensbachi-an Margaritatus Chronozone MNHNL-BEL256 BEL257 and BEL265

Description Small-sized more or less slender elongate and laterally compressed rostrum with an obtuse apex Some individuals bear a reduced mucro The outline is cy-lindrical The profile is asymmetrical subhastate with a more inflated venter in apical region The transverse sec-tion is subquadrate to elliptical in the apical region with a pronounced compression along the rostrum There are no grooves Lateral lines are more or less pronounced on the anterior part of the rostrum

Remarks See P aff compressus

Stratigraphic and geographic range Upper Pliensbachi-an England Germany France Luxembourg Switzerland Slovakia

Pleurobelus aff compressus (stahl 1824)Fig 5C D

1848 Belemnites acuarius amalthei ndash QuensteDt p 406 [pars] pl 24 figs 11-12

Material Seven rostra from the level 85 Upper Pliensbachi-an Margaritatus Chronozone MNHNL-BEL258 BEL259 BEL266 and BEL271

Description Small-sized more or less slender and elon-gate orthorostrum with an obtuse apex Some individuals bear a reduced mucro The outline is cylindrical The pro-file is asymmetrical subhastate with a more inflated venter in apical region The transverse section is subquadrate to elliptical in the apical region with a pronounced compres-sion along the orthorostrum There are no grooves on the orthorostrum Lateral lines are more or less pronounced on the anterior part of the orthorostrum The epirostrum is conical its outline and profile are symmetrical with a slightly compressed section Two dorso-lateral grooves and several striae are visible along the epirostrum The distinc-tion between the epirostrum and the orthorostrum is well-pronounced

Remarks The orthorostrum of P aff compressus is simi-lar to the rostrum of P compressus The only difference lies in the presence of an additional epirostrum in P aff com-pressus Both species are described as distinct morphospe-cies but could be considered as dimorphs from one biologi-cal species as stated by Doyle (1987) in the case of Cus-piteuthis trivialis (siMpson 1855) and C tubularis (young amp biRD 1922) P aff compressus differs from P lagenae-formis by the morphology of their respective orthorostrum

Stratigraphic and geographic range Upper Pliensbachi-an France Germany

Pleurobelus lagenaeformis (haRtMann in V Zieten 1832)

Fig 5E-G

1832 Belemnites lagenaeformis haRtMann ndash V Zieten p 33 pl 25 fig 1

1848 Belemnites acuarius amalthei ndash QuensteDt p 406 [pars] pl 24 figs 10 13-14

1998 Pleurobelus lagenaeformis (haRtMann) ndash schle-gelMilch p 54 pl 4 figs 4-6

Material Eighteen rostra from level 85 Upper Pliensbachi-an Margaritatus Chronozone MNHNL-BEL260 to 262 BEL267 and BEL272

Description Small-sized cylindriconical orthorostrum with slightly obtuse apex Some individuals bear a reduced mucro The outline and the profile are symmetrical and quite similar The transverse section is rounded to ellipti-cal and in the latter case slightly compressed There are no grooves on the orthorostrum The epirostrum is elongate and tubular Two dorso-lateral grooves are visible The dis-tinction between the epirostrum and the orthorostrum is well-pronounced

Remarks The orthorostrum of P lagenaeformis is similar to the rostrum of P subirregularis The only difference be-tween both is the presence of an epirostrum in P lagenae-formis Both species are described as distinct morphospe-cies but could be considered as dimorphs from one biologi-cal species (cf remark for P aff compressus)


Pleurobelus subirregularis (lissajous 1927)Fig 5H-I

1848 Belemnites acuarius amalthei ndash QuensteDt p 406 [pars] pl 24 fig 9

1927 Dactyloteuthis subirregularis sp n ndash lissajous p 18 pl 1 figs 10-12

1998 Pleurobelus subirregularis (lissajous) ndash schlegel-Milch p 55 pl 4 fig 7

Material Sixteen rostra from level 85 Upper Pliensbachi-an Gibbosus Subchronozone MNHNL-BEL263 BEL264 BEL268 and BEL270

Description Small-sized cylindriconical rostrum with slightly obtuse apex Some individuals bear a reduced mucro The outline and the profile are symmetrical and similar The transverse section is rounded to elliptical and in the latter case slightly compressed There are no grooves

Remarks See P lagenaeformis



Genus Rhabdobelus naef 1922

Type species Belemnites exilis DrsquooRbigny 1842 Toarcian iron ore of Saint Quentin-Fallavier (La Verpilliegravere) Isegravere France

Rhabdobelus donovani RiegRaf in RiegRaf et al 1998

Fig 5J-K

1856 Belemnites parvus haRtMann ndash QuensteDt p 286 pl 41 fig 21

1971 Belemnites parvus (haRtMann) ndash schWegleR p 115 text-figs 119-120

1998 Rhabdobelus donovani nom nov ndash RiegRaf et al p 85

Material One hundred and thirteen rostra from the level 104 to 106 Upper Toarcian Dispansum and Pseudoradiosa chronozones MNHNL-BEL299 to 309

Description Small-sized slender and elongate rostrum with an acute apex The outline is subhastate The profile is asymmetrical and hastate The transverse section is sub-quadrate in the anterior part of the rostrum and also ventral-ly flattened in the alveolar region A dorso-alveolar groove extends until the anterior part of the rostrum solidum This groove is more or less pronounced and sometimes replaced by a flat area Two well-developed lateral lines are visible on the rostrum cavum and extend on the two thirds of the anterior part of the rostrum solidum

Stratigraphic and geographic range Upper Toarcian France Germany

Rhabdobelus aff donovani RiegRaf in RiegRaf et al 1998

Fig 5L

Material Twenty-nine rostra from the level 100 to 104 Up-per Toarcian Thouarsense andor Dispansum chronozones MNHNL-BEL293 to 298 and BEL310

Description Small sized slender and elongate rostrum with an acute apex The outline is cylindriconical The pro-file is asymmetrical and slightly subhastate The transverse section is elliptical to slightly subquadrate Two lateral lines are visible on the rostrum cavum and extend on the first third of the anterior part of the rostrum solidum

Remarks This species is distinguished from R donovani by a short and conical rostrum slightly subhastate in some individuals The dorso-alveolar groove is always missing whereas well-developed in R donovani

Stratigraphic and geographic range Upper Toarcian Causses Basin France

Family Megateuthididae sachs amp nalnjaeVa 1967Genus Acrocoelites lissajous 1915

Type species Belemnites oxyconus hehl in Zieten 1831 Lower Toarcian Posidonienschiefer Formation of Bad Boll South-West Germany

Acrocoelites bobeti lissajous 1927Fig 6A

1927 Acrocoelites bobeti sp n ndash lissajous p 191971 Acrocoelites bobeti lissajous ndash coMbeacuteMoRel p 63

pl 1 figs 1-21990 Acrocoelites (Acrocoelites) bobeti lissajous ndash

Doyle p 33 pl 7 figs 9-11 13-15

Material Seventeen rostra from the 106 and 107 Upper Toarcian Pseudoradiosa Chronozone MNHNL-BEL311 to 316

Description Medium-sized slender and elongate rostrum with acute apex The outline and the profile are cylindrical in the stem and conical in the apical region However the profile is slightly asymmetrical and moderately subhastate near the alveolar part The transverse section is slightly compressed and elliptical near the alveolar part and round-ed otherwise Three apical grooves are visible two dorso-laterals and one longer ventral

Stratigraphic and geographic range Upper Toarcian England France Germany

Acrocoelites brevisulcatus (QuensteDt 1848)Fig 6B

1848 Belemnites acuarius brevisulcatus sp n ndash Quen-steDt p 413 pl 25 figs 1 3

1969 Belemnites acuarius brevisulcatus QuensteDt ndash schWegleR p 200 text-fig 83

1998 Acrocoelites (Odontobelus) brevisulcatus (Quen-steDt) ndash schlegelMilch p 62 pl 8 figs 4-5

Material Six rostra from the level 100 Upper Toarcian Thouarsense andor Dispansum chronozones MNHNL-BEL349 to 353

Description Small sized short conical orthorostrum with acute apex The outline and the profile are similar and conical with a slightly more inflated venter in the posterior part of the profile The transverse section is compressed elliptical to subquadrate Three apical grooves are vis-ible two dorso-laterals and a less pronounced ventral one The epirostrum is elongate conical and bears on its entire length three grooves the ventral one being more or less pro-nounced or absent in some individuals


Fig 6


Remarks The orthorostrum of A brevisulcatus is similar to the rostrum of A pyramidalis The only difference be-tween both is the presence of an epirostrum in A brevi-sulcatus Both species could be considered as two morpho-types of the same biological species

Stratigraphic and geographic range Upper Toarcian Germany France

Acrocoelites ilminstrensis (phillips 1867)Fig 6C

1848 Belemnites tripartitus paxillosus ndash QuensteDt p 420 pl 26 figs 25-26

1867 Belemnites ilminstrensis sp n ndash phillips p 64 pl 12 fig 30

1984 Acrocoelites (Acrocoelites) ilminstrensis (hehl) ndash RiegRaf et al p 155 pl10 figs 2-3 5 9

1990 Acrocoelites (Toarcibelus) ilminsterensis [sic] (phillips) ndash Doyle p 39 pl 1 figs 1-7

Material Three rostra from the level 92 Lower Toarcian Bifrons Chronozone MNHNL-BEL333 to 335

Description Medium to large sized more or less slender and elongate rostrum with acute apex The outline is cy-lindriconical The profile is conical to cylindriconical and symmetrical The transverse section elliptical and slightly compressed Three apical grooves are visible two dorso-laterals and a longer and more incised ventral one There are no true lateral lines but distinct lateral flattenings are visible on the anterior part of the rostrum

Remarks A ilminstrensis shows a striking similarity with A oxyconus (hehl in V Zieten 1831) All the transitional morphologies are visible in our material and also in the fig-ured specimens in literature (Doyle 1990 RiegRaf et al 1984) In our opinion these species are only based on the separation of two extreme morphotypes A oxyconus (Fig 6E) being slightly more compressed and conical than A il-

minstrensis (Fig 6 C) (see also RiegRaf et al 1984 155) A future revision will possibly elucidate this hypothesis

Stratigraphic and geographic range Lower Toarcian Bulgaria England France Germany Luxembourg

Acrocoelites levidensis (siMpson 1855)Fig 6D

1855 Belemnites levidensis sp n ndash siMpson p 201990 Acrocoelites (Odontobelus) levidensis (siMpson) ndash

Doyle p 45 pl 14 fig 8 pl 15 4-72003 Belemnites levidensis siMpson [Acrocoelites (Od)

levidensis (siMpson)] ndash Doyle p 175 fig 15e-f2003 Belemnites alveolatus siMpson [Acrocoelites (Od)

levidensis (siMpson)] ndash Doyle p 175 fig 12g-h

Material Nine rostra from the level 92 to 96 Lower Toar-cian Bifrons Chronozone MNHNL-BEL340 to 346

Description Medium- to large-sized and robust rostrum with acute apex The outline is cylindriconical The profile is conical and asymmetrical with a slightly inflated ventral part The transverse section is compressed and elliptical to slightly subquadrate Three apical grooves are visible two dorso-laterals and one ventral with the latter more pro-nounced and longer

Stratigraphic and geographic range Toarcian England France Germany Luxembourg

Acrocoelites oxyconus (hehl in V Zieten 1831)Fig 6E

1831 Belemnites oxyconus hehl ndash v Zieten p 27 pl 21 fig 5

1848 Belemnites tripartitus oxyconus sp n ndash QuensteDt p 419 pl 26 fig19

1984 Acrocoelites (Acrocoelites) oxyconus (hehl) ndash Rie-gRaf et al p 154 pl 10 figs 4 10

Fig 6 Belemnites from Tournadous Megateuthididae The specimens are all in natural size except where stated otherwise A ndash Acrocoelites bobeti lissajous 1927 BEL-311 rostrum in ventral view (A1) lateral view (A2) B ndash Acrocoelites brevi-sulcatus (QuensteDt 1848) BEL-349 rostrum in ventral view (B1) lateral view (B2) C ndash Acrocoelites ilminstrensis (phil-lips 1867) BEL-333 rostrum in ventral view (C1) lateral view (C2) D ndash Acrocoelites levidensis (siMpson 1855) BEL-346 rostrum in ventral view (D1) lateral view (D2) E ndash Acrocoelites oxyconus (hehl in V Zieten 1831) BEL-254 rostrum in ventral view (E1) lateral view (E2) F ndash Acrocoelites pyramidalis (Muumlnster in v Zieten 1831) BEL-239 rostrum in ven-tral view (F1) lateral view (F2) G ndash Acrocoelites pyramidalis (MuumlnsteR in V Zieten 1831) BEL-240 rostrum in ventral view (G1) lateral view (G2) H ndash Acrocoelites pyramidalis (MuumlnsteR in V Zieten 1831) BEL-241 rostrum in ventral view (H1) lateral view (H2) I ndash Acrocoelites conoideus (oppel 1856) BEL-326 rostrum in ventral view (I1) lateral view (I2) J ndash Acrocoelites cf riegrafi Doyle 1992 BEL-338 rostrum in ventral view (J1) lateral view (J2) K ndash Acrocoelites rostri-formis (theoDoRi in bRonn 1837) BEL-242 rostrum in ventral view (K1) lateral view (K2) L ndash Acrocoelites rostriformis (theoDoRi in bRonn 1837) BEL-243 rostrum in ventral view (L1) lateral view (L2)


1990 Acrocoelites (Acrocoelites) oxyconus (Zieten) ndash Doyle p 29 pl 6 figs1-4

Material Six rostra from the level 92 to 95 Lower Toar-cian Bifrons Chronozone MNHNL-BEL254 BEL255 and BEL142

Description Medium- to large-sized more or less slender and elongate rostrum with acute apex The outline is cy-lindriconical The profile is conical to moderately cylindri-conical and symmetrical The transverse section is elliptical and compressed Three apical grooves are visible two dor-so-laterals and a longer and more incised ventral one There are no true lateral lines only lateral flattenings are visible

Remarks See A ilminstrensis

Stratigraphic and geographic range Lower Toarcian England France Germany Slovakia

Acrocoelites pyramidalis (MuumlnsteR in V Zieten 1831)

Fig 6F-H

1831 Belemnites pyramidalis MuumlnsteR ndash v Zieten p 31 pl 24 fig 5

1998 Acrocoelites (Odontobelus) pyramidalis (Muumln-steR) ndash schlegelMilch p 62 pl 8 figs1-2

1998 Acrocoelites (Odontobelus) pyramidalis (Zieten) ndash coMbeacuteMoRel in Rulleau et al p 21 pl 1 fig 8

Material One hundred and nineteen rostra from the level 95 to 106 Lower to Upper Toarcian Bifrons to Pseudora-diosa chronozones MNHNL-BEL239 to 241 BEL244 to 250 and BEL330

Description Small-sized short conical rostrum with acute apex The outline and the profile are similar and conical with a slightly more inflated venter in the posterior part of the profile The transverse section is compressed elliptical to subquadrate Three apical grooves are visible two dorso-laterals and a less pronounced ventral one

Stratigraphic and geographic range Lower to Upper Toarcian England France Germany Luxembourg Slova-kia possibly Caucasus

Acrocoelites conoideus (oppel 1856)Fig 6I

1848 Belemnites compressus conicus sp n ndash QuensteDt p 423 pl 27 fig 4

1856 Belemnites conoideus sp n ndash oppel p 4831998 Acrocoelites (Acrocoelites) conoideus (oppel) ndash

schlegelMilch p 60 pl 6 figs7-8

Material One rostrum from the level 111 Upper Toarcien Mactra Subchronozone MNHNL-BEL326

Description Large and robust rostrum with moderately acute apex The outline is cylindriconical The profile is cylindriconical and asymmetrical with a slightly inflated ventral part The transverse section is rounded to slightly elliptical Two dorso-lateral and one ventral apical grooves are equally developed

Stratigraphic and geographic range Upper Toarcian to Lower Aalenian Bulgaria Caucasus France Germany Luxembourg Slovakia and possibly Romania

Acrocoelites cf riegrafi Doyle 1992Fig 6J

cf 1969 Belemnites longiconus sp n ndash schWegleR p 187 text-fig 73

cf 1990 Acrocoelites (Acrocoelites) longiconus (schWeg-leR) ndash Doyle p 34 pl 7 figs 4-6 17

cf 1992 Acrocoelites (Acrocoelites) riegrafi nom nov ndash Doyle p 75 [pro A (A) longiconus (schWegleR)]

cf 1998 Acrocoelites (Acrocoelites) riegrafi Doyle ndash schlegelMilch p 59 pl 5 fig 10

MaterialOne rostrum from the Lower Toarcian Bifrons Chronozone MNHNL-BEL338

DescriptionSmall-sized cylindriconical orthorostrum The outline and the profile are similar and cylindriconical The transverse section is compressed elliptical The epirostrum is short conical and bears two well-developed dorso-lateral grooves and one weakly developed ventral groove

Remarks Our sample shows a strong resemblance with the figured type specimen (schlegelMilch 1998 pl 5 fig 10) but differs by its reduced ventral groove

Stratigraphic and geographic range Lower Toarcian Causses Basin France (this report)

Acrocoelites rostriformis (theoDoRi in bRonn 1837)Fig 6K-L

1837 Belemnites rostriformis theoDoRi ndash bRonn p 4121848 Belemnites rostriformis theoDoRi ndash QuensteDt p

425 pl 27 figs 19-201998 Acrocoelites (Odontobelus) curtus (DrsquooRbigny) ndash

schlegelMilch p 62 pl 5 fig 14 pl 8 fig 31998 Acrocoelites curtus (DrsquooRbigny) ndash coMbeacuteMoRel in

Rulleau et al p 29 pl 2 figs 8-10

Material Nine rostra from the level 100 to 106 Upper Toarcian Thouarsense andor Dispansum to Pseudoradiosa chronozones MNHNL-BEL242 BEL243 BEL251 to 253 and BEL347

Description Small-sized short conical rostrum with an acute apex The outline is conical The profile is conical and asymmetrical with an inflated ventral part The trans-


verse section is rounded to elliptical in the anterior part and subpyriform in the apical region Three apical grooves are visible two well-developed dorso-lateral grooves and a ven-tral one reduced in some individuals

Stratigraphic and geographic range Upper Toarcian to lowermost Aalenian France Germany Luxembourg

Acrocoelites inaequistriatus (siMpson 1855)Fig 7A

1855 Belemnites inaequistriatus sp n ndash siMpson p 241990 Acrocoelites (Toarcibelus) inaequistriatus (siMp-

son) ndash Doyle p 41 pl 13 figs 1-5 pl 14 figs 1-3 7

1998 Acrocoelites (Toarcibelus) inaequistriatus (siMp-son) ndash Rulleau et al p 24 pl 2 fig 1

2003 Belemnites inaequistriatus siMpson [Acrocoelites (Praepachyteuthis) inaequistriatus (siMpson)] ndash Doyle p 173 fig 13a-b

Material Two rostra from the level 100 Upper Toarcian Thouarsense andor Dispansum chronozones MNHNL-BEL336

Description Medium- to large-sized conical rostrum with epirostrum The outline and the profile of the entire rostrum are similar and conical The transverse section is moderate-ly elliptical compressed to rounded Irregular longitudinal striae and three well-developed apical grooves (two dorso-lateral and one ventral) are visible on the epirostrum

Stratigraphic and geographic range Lower to Upper Toarcian England France Luxembourg

Acrocoelites subtenuis (siMpson 1855)Fig 7B

1855 Belemnites subtenuis sp n ndash siMpson p 261990 Acrocoelites (Acrocoelites) subtenuis (siMpson) ndash

Doyle p 30 pl 6 figs 5 7-14 pl 7 figs 1-3 text-fig 15

2003 Belemnites subtenuis siMpson [Acrocoelites (Acro-coelites) subtenuis (siMpson)] ndash Doyle p 169 fig 9c-f k-l

Material Six rostra from the level 91-92 Lower Toarcian Serpentinum to Bifrons chronozones MNHNL-BEL327 to 329

Description Medium-sized slender and elongate cylindri-conical rostrum The outline is cylindriconical and slightly subhastate The profile is symmetrical and cylindriconical The transverse section is slightly elliptical compressed near the alveolar part rounded in the medium part and slightly depressed subpyriform in the apical region The venter of the apical region is moderately flattened Three apical grooves are visible two short dorso-laterals and one longer ventral

Stratigraphic and geographic range Lower Toarcian England France Germany

Acrocoelites tripartitus (V schlotheiM 1820)Fig 7C

1820 Belemnites tripartitus sp n ndash v schlotheiM p 48 [pars]

1942 Mesoteuthis banzensis sp n ndash kolb p 163 pl 11 figs 1-2 7

1996b Acrocoelites (Acrocoelites) tripartitus (V schlothe-iM) ndash RiegRaf p 27 fig 24c

2000 Acrocoelites tripartitus (V schlotheiM) ndash RiegRaf p 287 figs 3-4 18

Material Twelve rostra from the level 100 to 109 Upper Toarcian Variabilis to Aalensis chronozones MNHNL-BEL322 to 325

Description Medium-sized conical to cylindriconical ros-trum with an acute apex The outline is conical The profile is conical to cylindriconical and asymmetric with a moder-ately inflated venter The transverse section is moderately elliptical compressed Three apical grooves two dorso-lat-erals and one longer and more incised ventral are visible

Remarks The present species can be distinguished from A conoideus and A quenstedti (oppel 1856) by its more elongate and conical apical region

Stratigraphic and geographic range Upper Toarcian France Germany Luxembourg possibly Ukraine and Cau-casus

Acrocoelites vulgaris (young amp biRD 1822)Fig 7D

1822 Belemnites vulgaris sp n ndash young amp biRD p 258 pl 14 fig 1

1867 Belemnites vulgaris young amp biRD ndash phillips p 73 pl 16 figs 40-41

1992 Acrocoelites (Odontobelus) vulgaris (young amp biRD) ndash Doyle p 44 pl 11 figs 6-8 pl 13 figs 6-7 pl 14 figs 4-6 pl 15 figs 2-3

Material Two rostra from the levels 95 and 98 Lower to Upper Toarcian Bifrons to Variabilis chronozones MNHNL-BEL331 BEL 32 and BEL339

Description Large-sized robust and cylindriconical ros-trum with a slightly recurved acute apex The outline is cy-lindriconical The profile is symmetrical and conical The transverse section is moderately compressed and elliptical Three apical grooves two dorso-laterals and one longer and more incised ventral are visible



Fig 7


Genus Cuspiteuthis abel 1916

Type species Belemnites acuarius V schlotheiM 1820 Lower Toarcian (Falciferum Subzone) Bituminous Shales Formation Saltwick Bay North Yorkshire

Cuspiteuthis acuaria (V schlotheiM 1820)Fig 8A

1820 Belemnites acuarius sp n ndash V schlotheiM p 46 [pars]

1848 Belemnites acuarius gigas sp n ndash QuensteDt p 412 pl 25 figs 6-7

1984 Youngibelus gigas (QuensteDt) ndash RiegRaf et al p 158 pl 11 figs 9-11

1998 Youngibelus ohmdenensis nom nov ndash schlegel-Milch p 63 pl 8 figs 10-11

2000 Cuspiteuthis acuaria (V schlotheiM) ndash RiegRaf p 292 figs 9-10 13-14

Material Five rostra from the level 95 Lower Toarcian Bi-frons Chronozone MNHNL-BEL92 BEL291 and BEL292

Description Medium-sized conical orthorostrum with a more or less acute apex and a well-developed epirostrum The outline and the profile of the orthorostrum are conical The transverse section is rounded to elliptical and slightly compressed No grooves are visible on the orthorostrum The epirostrum is very elongate and conical to tubular in the posterior part

Remarks C acuaria shows a striking similarity with C tubularis (young amp biRD 1822) All the transitional mor-phologies are visible in our material and also in the figured specimens in literature (Doyle 1992 RiegRaf et al 1984) In our opinion these species are certainly based on the separation of two extreme morphotypes C acuaria being slightly more robust and conical than C tubularis A future revision will possibly elucidate this hypothesis

Stratigraphic and geographic range Lower to lowermost Upper Toarcian France Germany Luxembourg

Genus Dactyloteuthis bayle 1878

Type species Belemnites irregularis V schlotheiM 1813 Upper Toarcian (Variabilis Zone) Jurensismergel Forma-tion Upper Franconia (South Germany) (fide RiegRaf 2000)

Dactyloteuthis incurvata (V Zieten 1831)Fig 7E

1831 Belemnites incurvatus sp n ndash V Zieten p 29 pl 22 fig 7a-e

1984 Dactyloteuthis (Dactyloteuthis) incurvata (V Zie-ten) ndash RiegRaf et al p 161 pl 12 fig10 text-fig 48r

1998 Belemnites pygmeus V Zieten [Dactyloteuthis] ndash RiegRaf et al p 217

1998 Dactyloteuthis incurvata (V Zieten) ndash schlegel-Milch p 66 pl 9 figs 12-13

Material Two rostra from the Lower Toarcian Bifrons Chronozone MNHNL-BEL280 and BEL281

Description Small-sized and robust rostrum with an acute apex The outline is cylindriconical to slightly subhastate The profile is symmetrical and cylindriconical The trans-verse section is well-compressed rounded subquadrate and slightly pyriform in the apical region Three apical grooves are visible the ventral one is well-developed contrary to the dorso-laterals which are more or less incised and do not reach the tip of the apex

Remarks RiegRaf et al (1998 78) stated that Belemnites incurvatus V Zieten 1831 is a primary homonym of Be-lemnites incurvatus Raspail 1829 and replace it by Belem-nites pygmeus V Zieten 1831 whose holotype has been re-studied by RiegRaf (in RiegRaf et al 1998 78) However Belemnites pygmeus is possibly a nomen nudum andor no-men oblitum (personal communication W RiegRaf 2003) and the better-known name Belemnites incurvatus is herein employed in anticipation of further clarification

Stratigraphic and geographic range Lower Toarcian Bulgaria France Germany Luxembourg

Fig 7 Belemnites from Tournadous Megateuthididae The specimens are all in natural size except where stated otherwise A ndash Acrocoelites inaequistriatus (siMpson 1855) BEL-336 rostrum in ventral view (A1) lateral view (A2) B ndash Acrocoe-lites subtenuis (siMpson 1855) BEL-327 rostrum in ventral view (B1) lateral view (B2) C ndash Acrocoelites tripartitus (V schlotheiM 1820) BEL-322 rostrum in ventral view (C1) lateral view (C2) D ndash Acrocoelites vulgaris (young amp biRD 1822) BEL-339 rostrum in ventral view (D1) lateral view (D2) E ndash Dactyloteuthis incurvata (V Zieten 1831) BEL-280 rostrum in ventral view (E1) lateral view (E2) F ndash Dactyloteuthis irregularis (V schlotheiM 1813) BEL-282 rostrum in ventral view (F1) lateral view (F2) G ndash Dactyloteuthis cf semistriata (MuumlnsteR 1830) BEL-286 rostrum in ventral view (G1) lateral view (G2) H ndash Dactyloteuthis similis (V seebach 1864) BEL-288 rostrum in ventral view (H1) lateral view (H2) I ndash Dactyloteuthis wrighti (oppel 1856) BEL-348 rostrum in ventral view (I1) lateral view (I2)


Dactyloteuthis irregularis (V schlotheiM 1813)Fig 7F

1813 Belemnites irregularis sp n ndash V schlotheiM p 70 pl 3 fig 2

1998 Dactyloteuthis irregularis (V schlotheiM) ndash schle-gelMilch p 65 pl 9 figs 1-4

2000 Dactyloteuthis irregularis (V schlotheiM) ndash Rie-gRaf p 293 figs 1-2 17

Material Ten rostra from the level 100 Upper Toarcian Thouarsense Chronozone andor Insigne Subchronozone MNHNL-BEL282 to 285

Description Medium-sized robust and digit-like rostrum with very obtuse apex The outline and the profile are cylin-drical to cylindriconical The transverse section is strongly compressed and elliptical The presence and the size of the apical ventral groove are variable

Stratigraphic and geographic range Upper Toarcian Bulgaria England France Germany Luxembourg Poland Slovakia

Dactyloteuthis cf semistriata (V MuumlnsteR 1830)Fig 7G

cf 1830 Belemnites semistriatus sp n ndash MuumlnsteR p 15 pl 2 fig 4

cf 1848 Belemnites acuarius ventricosus sp n ndash Quen-steDt p 411 pl 25 fig 8

cf 1984 Dactyloteuthis (Cuspiteuthis) semistriata (Muumln-steR) ndash RiegRaf et al p 163 pl 12 fig 4 12 text-fig 48u

cf 1994 Dactyloteuthis irregularis (schlotheiM) [Belem-nites acuarius] ndash fischeR p 9 pl 1 figs 4-5

Material One adult rostrum from the level 100 Upper Toarcian Thouarsense andor Dispansum chronozones MNHNL-BEL286

Description Small-sized and digit-like orthorostrum with very obtuse apex The outline and the profile are cylindrical to cylindriconical The transverse section is strongly com-

pressed and elliptical No apical grooves are visible on our specimen and the epirostrum is missing just the scar is vis-ible

Remarks The preservation of the unique specimen does not allow a definitive determination The characteristic epirostrum is not preserved only a scar indicates its pres-ence

Stratigraphic and geographic range Upper Toarcian France (this report)

Dactyloteuthis similis (V seebach 1864)Fig 7H

1864 Belemnites similis sp n ndash V seebach p 158 pl7 fig 6

1998 Dactyloteuthis irregularis (schlotheiM) ndash Rulleau et al p 25 pl 2 figs 2-3

1998 Dactyloteuthis similis (V seebach) ndash schlegel-Milch p 65 pl 9 figs 5-6

Material Five rostra from the level 100 Upper Toarcian Thouarsense andor Dispansum chronozones MNHNL-BEL287 to 289

Description Medium-sized slightly elongate and digit-like rostrum with very obtuse apex The outline and the profile are cylindrical to cylindriconical The transverse section is distinctly compressed and elliptical There is no apical groove

Stratigraphic and geographic range Upper Toarcian France Germany Luxembourg

Dactyloteuthis wrighti (oppel 1856)Fig 7I

1848 Belemnites digitalis tripartitus sp n ndash QuensteDt p 418 pl 26 figs 14 31

1856 Belemnites wrighti sp n ndash oppel p 2401990 Acrocoelites (Odontobelus) wrighti (oppel 1856) ndash

Doyle p 48 pl 1 figs 4-7

Fig 8 Belemnites from Tournadous Megateuthididae Passaloteuthididae and Salpingoteuthididae The specimens are all in natural size except where stated otherwise A ndash Cuspiteuthis acuaria (V schlotheiM 1820) BEL-92 rostrum in ventral view (A1) lateral view (A2) B ndash Parapassaloteuthis cf polita (siMpson 1866) BEL-278 rostrum in ventral view (B1) lateral view (B2) C ndash Passaloteuthis bruguieriana (DrsquooRbigny 1842) BEL-275 rostrum in ventral view (C1) lateral view (C2) D ndash Passaloteuthis laevigata (V Zieten 1831) BEL-273 rostrum in ventral view (D1) lateral view (D2) E ndash Passalo-teuthis milleri (phillips 1867) BEL-277 rostrum in ventral view (E1) lateral view (E2) F ndash Pseudohastites sp BEL-277 rostrum in ventral view (F1) lateral view (F2) G ndash Salpingoteuthis dorsetensis (oppel 1856) BEL-357 rostrum in ventral view (G1) lateral view (G2) and dorsal view (G3) H ndash Salpingoteuthis trisulcata (De blainVille 1827) BEL-317 rostrum in ventral view (H1) lateral view (H2) and dorsal view (H3)


Fig 8


1994 Dactyloteuthis incurvata (Zieten) [Belemnites no-dotianus (DrsquooRbigny)] ndash fischeR p 13 pl 2 figs 5-6

Material Two rostra from the Lower Toarcian Bifrons Chronozone MNHNL-BEL348

Description Medium-sized and elongate rostrum with acute apex The outline and the profile are cylindriconical The transverse section is compressed and elliptical There is one short more or less incised ventral groove

Stratigraphic and geographic range Toarcian England France Germany Greenland Luxembourg

Family Passaloteuthididae naef 1922Genus Parapassaloteuthis RiegRaf 1980 (emend

Doyle 1990)

Type species Belemnites zieteni WeRneR 1912 [= Belem-nites zieteni MayeR-eyMaR 1884 fide RiegRaf et al 1998] Upper Pliensbachian Amaltheenton Formation South-west Germany

Parapassaloteuthis cf polita (siMpson 1866)Fig 8B

cf 1866 Belemnites politus sp n ndash siMpson p 216cf 1990 Parapassaloteuthis polita (siMpson) ndash Doyle p

27 pl 5 figs 3-9cf 2003 Belemnites politus siMpson [Parapassaloteuthis

polita (siMpson)] ndash Doyle p 167 fig 6m-n

Material One rostrum from the level 85 Upper Pliensba-chian Margaritatus Chronozone MNHNL-BEL278

Description Small-sized robust rostrum with a slightly mucronate apex The outline is cylindriconical The pro-file is cylindriconical and asymmetrical with more inflated venter The transverse section is slightly compressed and rounded subquadrate Two dorso-lateral apical grooves are weakly developed

Remarks Our specimen shares morphological charac-ters of Pa brevis (De blainVille 1827) from the Lower Pliensbachian and Pa robusta (siMpson 1855) from the Lower Toarcian It resembles most closely some Pa polita in Doyle (1990 pl 5 figs 6-7) but lacks the typical subh-astate outline as given in the diagnosis (Doyle 1990 27)

Stratigraphic and geographic range Upper Pliensbach-ian France (this report)

Genus Passaloteuthis lissajous 1915

Type species Belemnites bruguierianus DrsquooRbigny 1842 Pliensbachian France

Passaloteuthis bruguieriana (DrsquooRbigny 1842)Fig 8C

1842 Belemnites bruguierianus sp n ndash DrsquooRbigny p 841990 Passaloteuthis bisulcata (blainVille) [variety A] ndash

Doyle p 19 pl 1 figs 1-2 pl 2 fig 11990 Passaloteuthis bisulcata (blainVille) [variety B] ndash

Doyle p 19 pl 2 figs 2-31998 Passaloteuthis bisulcata (blainVille) ndash schlegel-

Milch p 51 pl 2 figs 6-7

Material Two rostra from the level 85 Upper Pliensbachi-an Margaritatus Chronozone MNHNL-BEL275 and BEL276

Description Large-sized robust cylindriconical rostrum with a more or less obtuse apex The outline and the profile are cylindriconical The transverse section is rounded Two weakly developed dorso-lateral apical grooves are present

Stratigraphic and geographic range Upper Pliensbachian to lowermost Toarcian Europe (Bulgaria England France Germany Italy Luxembourg Portugal Slovakia Spain) northern Africa (Algeria Morocco) Western Asia (Turkey)

Passaloteuthis laevigata (V Zieten 1831)Fig 8D

1831 Belemnites laevigatus sp n ndash v Zieten p 28 pl 21 fig 12

1990 Passaloteuthis bisulcata (blainVille) [variety C] ndash Doyle p 19 pl 3 figs 1-3

1998 Passaloteuthis laevigata (Zieten) ndash schlegelMilch p 51 pl 2 fig 8

Material Six rostra from the level 85 Upper Pliensbachian Margaritatus Chronozone MNHNL-BEL273 and BEL274

Description Large-sized elongate cylindriconical rostrum with a moderately acute apex The outline and the profile are cylindriconical to slightly conical The transverse sec-tion is rounded subquadrate Two well-developed dorso-lateral apical grooves are present

Stratigraphic and geographic range Upper Pliensbachi-an to lowermost Toarcian England France Germany Lux-embourg

Passaloteuthis milleri (phillips 1867)Fig 8E

1867 Belemnites milleri sp n ndash phillips p 54 pl 8 fig 19

1992 Passaloteuthis milleri (phillips) ndash Doyle p 22 pl 2 figs 5-9

1998 Passaloteuthis milleri (phillips) ndash schlegelMilch p 52 pl 3 fig 4 pl 7 fig 6


Material Two rostra from the level 90 Upper Pliensbach-ian MNHNL-BEL277

Description Small-sized cylindrical rostrum with an ob-tuse apex The outline is cylindrical and the profile cylindri-conical The transverse section is rounded to subpyriform near the apex No apical grooves are visible

Stratigraphic and geographic range Upper Pliensbach-ian to lowermost Toarcian Bulgaria England France Ger-many Luxembourg

Genus Pseudohastites naef 1922

Type species Belemnites scabrosus siMpson 1866 Lower Pliensbachian (Jamesoni Zone) Robin Hoodrsquos Bay North Yorkshire

Pseudohastites spFig 8F

Material One adult rostrum from the level 85 Up-per Pliensbachian Margaritatus Chronozone MNHNL-BEL337

Description Large-sized slender and elongate orthoros-trum with a moderately acute apex The outline and the profile are similar and subhastate The transverse section is rounded subquadrate in the anterior part of the rostrum to rounded in the posterior part A strong dorso-alveolar flat-tening is present The epirostrum is conical to subtubular and bears two shallow dorso-lateral grooves

Remarks The specimen fits well the diagnosis of Pseu-dohastites (Doyle 1990 23) but cannot be referred to any known species for its peculiar characters the strong dorso-alveolar flattening and a true epirostral development

Stratigraphic and geographic range Upper Pliensbachi-an France (this report)

Family Salpingoteuthididae Doyle 1992Genus Salpingoteuthis lissajous 1915

Type species Belemnites trisulcatus De blainVille 1827 Toarcian Calvados Northwest France

Salpingoteuthis dorsetensis (oppel 1856)Fig 8G

1856 Belemnites dorsetensis sp n ndash oppel p 3621992 Salpingotheuthis dorsetensis (oppel) ndash Doyle p

72 pl 27 figs 1-21998 Salpingotheuthis dorsetensis (oppel) ndash Rulleau et

al p 28 pl 2 fig 7

Material Two rostra from the level 106 Upper Toar-cian Pseudoradiosa Chronozone MNHNL-BEL357 and BEL358

Description Short reduced conical orthorostrum with acute apex The outline is conical The profile is asymmet-rical and conical The transverse section is rounded The epirostrum is strongly developed (90 of the total rostrum) and conical to short tubular A broad apical ventral groove several apical striae and lateral lines fading out near the apex are visible

Stratigraphic and geographic range Upper Toarcian to Aalenian Bulgaria Caucasus England France Germany

Salpingoteuthis trisulcata (De blainVille 1827)Fig 8H

1827 Belemnites trisulcatus sp n ndash De blainVille p 83 pl 5 fig 13

1848 Belemnites acuarius tricanaliculatus sp n ndash Quen-steDt p 414 pl 25 figs 13-14

1992 Salpingoteuthis trisulcata (blainVille) ndash Doyle p 71 pl 28 figs 2-5 10

1994 Salpingoteuthis trisulcata (blainVille) ndash fischeR p 13 pl 2 figs 12-13

Material Twelve rostra from the level 109 and 110 Upper Toarcian Aalensis Chronozone MNHNL-BEL317 to 321

Description Short reduced conical orthorostrum with acute apex The profile is conical The transverse section is rounded The epirostrum is strongly developed (90 of the total rostrum) and very elongate tubular The dorso-lateral grooves paired in the anterior part are always well-developed The incision of the dorsal and ventral grooves extending to the alveolar region is highly variable Several irregular striae are present

Fig 9 Specific richness throughout the studied period The condensate level is represented by a single bar for the Thouarsense Chronozone and the beginning of the Dispan-sum Chronozone by the fact of the melting of both faunas (Ma Margaritatus Sp Spinatum Te Tenuicostatum Ser-pentinum Bi Bifrons Va Variabilis Th Thouarsense Di Dispansum Ps Pseudoradiosa Aa Aalensis)


Stratigraphic and geographic range Upper Toarcian to Aalenian England France Germany Slovakia

5 Discussion

The analysis of the belemnite fauna reveals that specif-ic richness is not constant over the studied period (Fig 9) Due to lack of data the diversity variations during the Late Pliensbachian cannot be described precisely However it is possible to observe an important spe-cies richness decrease between the Margaritatus and Serpentinum chronozones From the Serpentinum to Bifrons chronozones species richness rapidly rises reaching a maximum during the Bifrons Chronozone Subsequently the species richness strongly decreases (in Variabilis Chronozone) and then recovers reach-ing a peak during Thouarsense Chronozone (or the beginning of Dispansum Chronozone) with similar levels than previously seen in the Bifrons Chrono-zone It is however not possible to be more precise on the position of this peak due to a condensation of the Thouarsense Chronozone and the beginning of the Dispansum Chronozone During the late Dispansum Chronozone species richness slightly decreases and remains stable during the late Toarcian Pseudoradiosa and Aalensis chronozones share an average level spe-cies richness compared to Toarcian variations

The change in specific richness between Margari-tatus and Serpentinum chronozones is certainly related to the Toarcian Oceanic Anoxic Event occurring dur-ing the Early Toarcian Crisis The post-crisis recovery of high specific richness in belemnites occurring dur-ing Bifrons Chronozone is a common pattern already observed in belemnites (little amp benton 1995) and also in ammonites (DeRa et al 2010)

During the studied period it is also possible to observe variations at the generic scale The most rel-evant turnover concerns the Pliensbachian and the earliest Toarcian with the disappearance of all the Pliensbachian genera (Passaloteuthis Parapassalo-teuthis Pseudohastites and Pleurobelus) and the rise of an Acrocoelites-dominated population During the late Toarcian the faunal renewal is less sudden with a progressive demise of early Toarcian genera (Acro-coelites Cuspiteuthis and Dactyloteuthis) and the ap-pearance and domination of new taxa (Rhabdobelus Salpingoteuthis)

The faunal composition in Tournadous is relative-ly different from those known from England (Doyle 1990 1992) and South-west Germany (RiegRaf 1980 RiegRaf et al1984) In Tournadous the genera Mega-teuthis and Brevibelus are absent whereas they are

rather abundant in the northern part of the Northwest European Province (England and Southern Germany) A similar pattern is observed for Southern France in the Aalenian-Bajocian (see MaRiotti et al 2012) In the same way there are relatively few Dactyloteuthis in Tournadous On the contrary there is a predomi-nance of the genera Rhabdobelus and Salpingoteuthis in the late Toarcian a pattern which is quite differ-ent from the northern part of the Northwest European Province (Doyle 1992 Weis 1999) These apparent absencespredominances of the mentioned genera hint at an onsetting provincialism during the Toarcian with ldquosouthern or Tethyan belemnite faunasrdquo as sug-gested by Doyle (1987 1994) and subsequently for the Aalenian by MaRiotti et al (2012)

Based on the study of a single outcrop it is impos-sible at the present day to make a detailed compari-son with the well-known faunas of the northern part of the Northwest European Province (such as done for the same geological ages by DeRa et al 2011 for ammonites and based on a very complete database) Thus the observed diversity and palaeobiogeographic patterns could be only local patterns (ie taphonomic or environmental effects) and not extensible to other palaeogeographic areas of the southern part of the Northwest European Province However the observed patterns are coherent with previous observations of other studies (Doyle 1987 1994 MaRiotti et al 2012) which indicate slight differences in faunal composition between the northern parts of the Northwest European Province on the one hand and the southern parts of the Northwest European Province plus the Mediterranean Province on the other In order to confirm these obser-vations it will then be necessary to extend this kind of study to further localities in the same palaeogeograph-ic region or others in the Mediterranean Province to gain a better understanding of the palaeobiogeography of the belemnites

Acknowledgements

We thank chRistian MeisteR (Museum of Natural His-tory Geneva Switzerland) for his help and the sharing of his knowledge on the biostratigraphy of the studied area The field work has been possible only by the commitment of guy and liette kRonZ-WilMes (Luxembourg) RobeRt haas (Luxembourg) and alain fabeR (National Museum of Natural History Luxembourg) DoMenico Mannetta (Sapienza University Rome Italy) prepared the figured thin sections Our acknowledgements go also to oksana DZyuba and peteR Doyle for their advices during the review pro-cess which allowed to improve our manuscript This work is a contribution by the INTERVIE (INSU) research pro-


gram and by the BioME team of the Biogeacuteosciences labora-tory (Dijon France) The work of jean-Daniel pinaRD is supported by the Fonds National de la Recherche (Luxem-bourg)

References

batheR fa (1888) Professor Blake and shell-growth in Cephalopoda ndash Annals and Magazine of Natural His-tory series 6 1 421-427

bayle E (1878) Fossiles principaux des terrains ndash Expli-cation de la carte geacuteologique de France 4 1-158 Paris (Imprimerie Nationale)

blainVille MH DucRotay De (1827) Meacutemoire sur les beacutelemnites consideacutereacutees zoologiquement et geacuteologique-ment ndash 136 pp Paris (Levrault)

bRonn HG (1835-1837) Lethaea geognostica oder Abbil-dungen und Beschreibungen der fuumlr die Gebirgs-Forma-tionen bezeichnendsten Versteinerungen Erster Band das Uumlbergangs- bis Oolithengebirge beinhaltend ndash 544 pp Stuttgart (Schweizerbart)

coMbeacuteMoRel R (1971) Figuration et compleacutement de dia-gnose de Acrocoelites bobeti lissajous beacutelemnite du Lias supeacuterieur franccedilais ndash Geobios 4 61-68

DeRa G neige P DoMMeRgues JL faRa E laffont R amp pellenaRD P (2010) High-resolution dynam-ics of Early Jurassic marine extinctions the case of PliensbachianndashToarcian ammonites (Cephalopoda) ndash Journal of the Geological Society (London) 167 21-33

DeRa G neige P DoMMeRgues JL amp bRayaRD A (2011) Ammonite paleobiogeography during the Pliensbachian-Toarcian crisis (Early Jurassic) reflect-ing paleoclimate eustasy and extinctions ndash Global and Planetary Change 78 92-105

DoMMeRgues JL MeisteR C amp MouteRDe R (1997) Pliensbachian ndash In caRiou Eacute amp hantZpeRgue P (Eds) Biostratigraphie du Jurassique ouest-europeacuteen et meacutediterraneacuteen zonations parallegraveles et distribution des inverteacutebreacutes et microfossiles ndash Bulletin du Centre de Re-cherches Elf Exploration et Production Meacutemoires 17 15-24

Doyle P (1987) Lower Jurassic-Lower Cretaceous belem-nite biogeography and the development of the Mesozoic Boreal Realm ndash Palaeogeography Palaeoclimatology Palaeoecology 61 237-254

Doyle P (1990) The British Toarcian (Lower Jurassic) be-lemnites Part 1 ndash Monographs of the Palaeontographi-cal Society 144 1-49


Doyle P (1994) Aspects of the distribution of Early Jurassic belemnites ndash Proceedings of the Third Pergola International Symposium ndash Palaeopelagos Special Publications 1 109-120

Doyle P (2003) Type belemnites of Simpsonrsquos fossils of the Yorkshire Lias ndash Proceedings of the Yorkshire Geo-logical Society 54 147-184

Doyle P amp kelly SRA (1988) The Jurassic and Creta-ceous belemnites of Kong Karls Land ndash Norsk-Polarin-

stitutt Skrifter 189 1-77elMi S Rulleau L gabilly J amp MouteRDe R (1997)

Toarcien ndash In caRiou Eacute amp hantZpeRgue P (Eds) Biostratigraphie du Jurassique ouest-europeacuteen et meacutedi-terraneacuteen zonations parallegraveles et distribution des in-verteacutebreacutes et microfossiles ndash Bulletin du Centre de Re-cherches Elf Exploration et Production Meacutemoires 17 25-36

fischeR JC (1994) Reacutevision critique de la Paleacuteontologie Franccedilaise drsquoAlcide drsquoOrbigny 1 8-22 Paris (Museacuteum national drsquoHistoire naturelle)

gRaciansky PC DaRDeau G DoMMeRgues JL DuR-let C MaRchanD D DuMont T hesselbo SP jacQuin T goggin V MeisteR C MouteRDe R Rey J amp Vail P (1998) Ammonite biostratigraphic correlation and Early Jurassic sequence stratigraphy in France comparisons with some UK sections ndash In De gRaciansky P-C et al (Eds) Mesozoic and Cenozoic sequencestratigraphy of European basins ndash SEPM Spe-cial Publications 60 583-621

guex J (1972) Reacutepartition biostratigraphique des ammo-nites du Toarcien moyen de la bordure sud des Causses (France) et reacutevision des ammonites deacutecrites et figures par MonestieR (1931) ndash Eclogae Geologicae Helvetiae 65 611-645

guex J MoRaRD A baRtolini A amp MoRettini E (2001) Discovery of an important stratigraphic gap at the Domerian-Toarcian limit palaeo-oceanographic implications ndash Bulletin de la Socieacuteteacute vaudoise des sci-ences naturelles 87 277-284

iba Y sano S MutteRlose J amp konDo Y (2012) Be-lemnites originated in the Triassic ndash A new look at an old group ndash Geology 40 (10) 911-914

kolb H (1942) Die Belemniten des juumlngeren Lias zeta in Nord Bayern ndash Zeitschrift der Deutschen geologischen Gesellschaft 94 145-168

lissajous M (1915) Quelques remarques sur les beacutelemni-tes jurassiques ndash Bulletin de la Socieacuteteacute drsquoHistoire natu-relle (Macirccon) 6 1-32

lissajous M (1925) Reacutepertoire alphabeacutetique des beacutelem-nites jurassiques preacuteceacutedeacute drsquoun essai de classification ndash Travaux du Laboratoire de Geacuteologie de la Faculteacute des Sciences de Lyon 8 (7) 1-173

lissajous M (1927) Description des quelques nouvelles espegraveces de beacutelemnites jurassiques ndash Travaux du Labo-ratoire de Geacuteologie de la Faculteacute des Sciences de Lyon 10 (7) 1-42

little CTS amp benton MJ (1995) Early Jurassic mass extinction A global long-term event ndash Geology 23 (6) 495-498

Maillot S Mattioli E baRtolini A bauDin F pit-tet B amp guex J (2009) Late Pliensbachian-Early Toarcian (Early Jurassic) environmental changes in an epicontinental basin of NW Europe (Causses area cen-tral France) A micropaleontological and geochemical approach ndash Palaeogeography Palaeoclimatology Pal-aeoecology 273 346-364

MaRiotti N Weis R Di cencio A cleacuteMent A amp De baets K (2012) New records of early Middle Jurassic belemnites in the French Subalpine Basin and their paleobiogographic significance ndash Geobios 45 99-108


MayeR-eyMaR k (1884) Die Filiation der Belemnites acu-ti ndash Vierteljahresschriften der Naturforschenden Ge-sellschaft in Zuumlrich 29 41-56

MeisteR C (1989) Les ammonites du Domeacuterien des Caus-ses ndash Cahiers de Paleacuteontologie ndash 80 pp Paris (Edition du CNRS)

MonestieR J (1921) Le Toarcien supeacuterieur de la reacutegion Sud-Est de lrsquoAveyron ndash Bulletin de la Socieacuteteacute geacuteologi-que de France 4 280-312

MonestieR J (1931) Ammonites rares ou peu connues et ammonites nouvelles du toarcien moyen de la reacutegion sud-est de llsquoAveyron ndash Meacutemoires de la Socieacuteteacute geacuteologi-que de France nouvelle seacuterie 15 1-70

MoRaRD A (2004) Les eacuteveacutenements du passage Domeacuterien-Toarcien entre Teacutethys occidentale et Eurpe du Nord-Ouest ndash Unpublished PhD Thesis Universiteacute de Lau-sanne ndash 417 pp

MuumlnsteR G gRaf Zu (1830) Bemerkungen zur naumlheren Kenntnis der Belemniten ndash 18 pp Bayreuth

naef A (1922) Die fossilen Tintenfische ndash 322 pp Jena (G Fischer)

oppel A (1856) Die Juraformation Englands Frankreichs und des suumldwestlichen Deutschlands nach ihren einzel-nen Gliedern eingetheilt und verglichen ndash Jahreshefte des Vereins fuumlr Vaterlaumlndische Naturkunde in Wuumlrttem-berg 12 121-556

oRbigny A De (1842) Paleacuteontologie Franccedilaise Terrains Ju-rassiques Tome I Ceacutephalopodes ndash 119 pp Paris (Mas-son)

page KN (2003) The Lower Jurassic of Europe its sub-division and correlation ndash In ineson JR amp suRlyk F (Eds) The Jurassic of Denmark and Greenland ndash Geo-logical Survey of Denmark and Greenland Bulletins 1 23-59

phillips J (1867) A monograph of British Jurassic Belem-nitidae ndash Monographs of the Palaeontographical Soci-ety 3 53-88

QuensteDt FA (1848) Petrefactenkunde Deutschlands 1 Die Cephalopoden 4 265-408

QuensteDt FA (1856-1857) Der Jura 1 1-368 Tuumlbingen (Laupp)

Raspail FV (1829) Histoire naturelle des beacutelemnites ac-compagneacutee de la description et de la classification des espegraveces que M Eacutemeric de Castellane a recueillies dans les Basses-Alpes de Provence ndash Annales des Sciences drsquoObservation 1 271-331

RiegRaf W (1980) Revision der Belemniten des Schwaumlbi-schen Jura Teil 7 ndash Palaeontographica (A) 169 128-206

RiegRaf W (2000) The belemnites described by Baron Ernst Friedrich von Schlotheim (1764-1833) ndash Palaumlon-tologische Zeitschrift 74 281-303

RiegRaf W (1996a) Belemniten-Fanghaumlkchen (Cephalo-poda Coleoidea) aus der Psilonotenbank (Unterer Jura tiefstes Hettangium) von Suumlddeutschland ndash Stuttgarter Beitraumlge zur Naturkunde (B) 239 1-38

RiegRaf W (1996b) Belemniten im Ober-Toarcium und Unter-Aalenium Suumldwestdeutschlands ndash In ohMeRt W et al Die Grenzziehung Unter-Mitteljura bei Wittnau und Fuentelsaz ndash Informationen des Geologischen Lan-desamtes Baden-Wuumlrttemberg 8 26-29

RiegRaf W WeRneR G amp loumlRcheR F (1984) Der Posi-donienschiefer Biostratigraphie Fauna und Fazies des suumldwestdeutschen Untertoarciums (Lias epsilon) ndash 195 pp Stuttgart (Enke)

RiegRaf W janssen N amp schMitt-RiegRaf C (1998) Cephalopoda dibranchiata fossiles (Coleoidea) II ndash In Westphal F (Ed) Fossilium Catalogus Animalia 135 1-519 Leiden (Backhuys)

Rulleau L alMeRas Y coMbeacuteMoRel R elMi S amp tin-tant H (1998) Critical revision of the cephalopods and brachiopods described by Eugegravene Dumortier (1874) in the tome IV (Toarcian Aalenian) of the ldquoEtudes paleacuteon-tologiques des deacutepocircts jurassiques du Bassin du Rhocircnerdquo ndash Meacutemoires du Museacuteum drsquoHistoire naturelle de Lyon 2 18-30

sachs Vn amp nalrsquonyaeVa ti (1967) Contribution to the systematics of Jurassic and Cretaceous belemnites ndash In sachs VN (Ed) Problems of paleontologic substantia-tion of detailed Mesozoic stratigraphy of Siberia and the Far East USSR 2 6-26 Leningrad (NAUKA)

sanDeRs MT baRDin J benZaggagh M amp cecca F (2013) Early Toarcian (Jurassic) belemnites from north-eastern Gondwana (South Riffian ridges Morocco) ndash Palaumlontologische Zeitschrift DOI 101007s12542-013-0214-0

schlegelMilch R (1998) Die Belemniten des suumlddeut-schen Jura ndash 151 pp Stuttgart Jena Luumlbeck amp Ulm (G Fischer)

schlotheiM EF V (1813) Beitraumlge zur Naturgeschich-te der Versteinerungen in geognostischer Hinsicht ndash Leonhards Taschenbuch fuumlr die gesammte Mineralogie 7 3-134

schlotheiM EF V (1820) Die Petrefactenkunde auf ih-rem jetzigen Standpunkte durch die Beschreibung sei-ner Sammlung versteinerter und fossiler Uumlberreste des Thier- und Pflanzenreichs der Vorwelt erlaumlutert ndash 88 pp Gotha (Becker)

schWegleR E (1969) Revision der Belemniten des Schwauml-bischen Jura Teil 5 ndash Palaeontographica (A) 132 179-219


sciau J (1993) Coup drsquooeil sur les fossiles des Causses II Jurassique du Toarcien au Kimmeacuteridgien ndash 95 pp Millau (Association des amis du museacutee de Millau)

sciau J (2011) Fossiles des Schistes Carton du Toarcien infeacuterieur des Causses ndash 32 pp Millau (Association Pa-leacuteontologique des Causses)

seebach K V (1864) Der Hannoversche Jura ndash 158 pp Berlin (Hertz)

siMpson M (1855) The fossils of the Yorkshire Lias de-scribed from nature 1st edition ndash 256 pp London amp Whitby (Wheldon)

siMpson M (1866) Inferior Oolite-Lias belemnites of the Yorkshire coast ndash Geological and Natural History Rep-ertory and Journal of Prehistoric Archaeology and Eth-nology 215-216

stahl CF (1824) Uumlbersicht uumlber die Versteinerungen Wuumlrttembergs ndash Korrespondenzblaumltter des Wuumlrttem-bergischen Landwirthschaftlichen Vereins 6 1-91


tRuumlMpy DM (1983) Le Lias moyen et supeacuterieur des Grands Causses et de la reacutegion de Rodez contributions stratigraphiques seacutedimentologiques et geacuteochimiques agrave la connaissance drsquoun bassin agrave seacutedimentation marneuse ndash Cahiers de lrsquoUniversiteacute Universiteacute de Pau et des Pays de lrsquoAdour 19 1-363

Van De schootbRugge B bachan A suan G RichoZ S amp payne JL (2013) Microbes mud and methane cause and consequence of recurrent early Jurassic an-oxia following the end-triassic mass extinction ndash Palae-ontology 56 685-709

Weis R (1999) die Belemniten der Minette-Formation (ob Toarcium-ob Aalenium) Luxemburgs ndash In Delsate D Duffin C amp Weis R Les collections paleacuteontologiques du Museacutee national drsquohistoire naturelle de Luxembourg Fossiles du Trias et du Jurassique ndash Travaux scientifi-ques du Museacutee national drsquohistoire naturelle de Luxem-bourg 32 201-246

Weis R amp Delsate D (2006) The earliest belemnites New records from the Hettangian of Belgium and Lux-embourg ndash In kostak M amp MaRek J (Eds) Pro-ceedings of the 2nd International Symposium Coleoid Cephalopods through time Prague 26-29 September 2005 ndash Acta Universitas Carolinae Geologica 49 181-184

Weis R amp MaRiotti N (2007) A belemnite fauna from the Aalenian-Bajocian boundary beds of the Grand Duchy of Luxembourg (NE Paris Basin) ndash Bollettino delle Societagrave Paleontologica Italiana 46 (2-3) 149-174

Weis R MaRiotti N amp RiegRaf W (2012) The belem-nite family Holcobelidae (Coleoidea) in the European Jurassic systematics biostratigraphy palaeobiogeogra-phy and evolutionary trends ndash Palaeodiversity 5 13-49

WeRneR e (1912) Uber die Belemniten des schwaumlbischen Lias und die mit ihnen verwandten Formen des Braunen Jura (Acoeli) ndash Palaeontographica 59 105-146

young G amp biRD J (1822) A geological survey of the Yorkshire Coast 1st edition ndash 335 pp Whitby (Clark)

Zieten KH V (1830-1833) Die Versteinerungen Wuumlrttem-bergs ndash 120 pp Stuttgart (Schweizerbart)

Zittel ka V (1895) Grundzuumlge der Palaumlontologie (Palaumlozoologie) ndash VIII + 971 pp Muumlnchen amp Leipzig (Oldenbourg)

Manuscript received February 10th 2014Revised version accepted by the Stuttgart editor June 13th 2014

Addresses of the authors

jean-Daniel pinaRD Scientific Research Centre of the Na-tional Museum for Natural History 25 rue Muumlnster 2160 Luxembourg Luxembourg and Biogeosciences Laboratory UMR6282 University of Burgundy CNRS 6 boulevard Gabriel 21000 Dijon Francee-mail jdpinardmnhnluRobeRt Weis anDRea Di cencio Scientific Research Cen-tre of the National Museum for Natural History 25 rue Muumlnster 2160 Luxembourg Luxembourge-mails rweismnhnlu andreadicenciogmailcompascal neige Biogeosciences Laboratory UMR6282 Uni-versity of Burgundy CNRS 6 boulevard Gabriel 21000 Dijon Francee-mail pascalneigeu-bourgognefrnino MaRiotti Earth Science Department Sapienza Uni-versity Piazzale A Moro 5 00185 Rome Italye-mail ninomariottiuniroma1it


of data is due to two factors (1) the low number of belemnites study in these areas and (2) some outcrops (ie in Italy and Spain) are hardened carbonates lead-ing a very difficult extraction of well-preserved speci-mens It is therefore impossible at the present day to portray the Jurassic belemnite diversification at the species level mostly because we lack data from some palaeogeographic areas unless introducing strong pal-aeogeographic biases

In the present study we report a belemnite fauna of more than 700 specimens from the Tournadous section in the Causses Basin This fauna constitutes then a homogeneous and comprehensive record of the Late Pliensbachian and Toarcian belemnite fauna in the southern part of the Northwest European Prov-ince These specimens have been collected in a com-mon field study associating ammonites and belemnites scholars It allows to assign the stratigraphic distribu-

tion of belemnites to a standard chronostratigraphic frame based on ammonites The reported belemnites substantially increase the knowledge of Early Jurassic faunas from the southern part of the Northwest Euro-pean Province a step that could later lead to further palaeogeographic explorations of the Jurassic radia-tion

2 Geological and palaeogeographic settings

We study here a single section (Tournadous) which ex-poses Lower Jurassic marls located in Central-South France (Fig 1) This section has been studied by sev-eral authors principally for its ammonoids content (eg MonestieR 1921 1931 guex 1972 MeisteR 1989 MoRaRD 2004) or to explore environmental changes at the Upper Pliensbachian-Lower Toarcian bound-

Fig 1 Palaeogeographic and geographic setting of Tournadous (Causses France) The provinces boundary is determined on ammonite fauna (Palaeogeographic map modified from DeRa et al 2010)









































Fig 5E-G




















Fig 5J-K








Fig 5L










Doyle p 33 pl 7 figs 9-11 13-15











Fig 6


































Fig 6F-H

































































Fig 7

























































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References






















































































































Fig 5E-G




















Fig 5J-K








Fig 5L










Doyle p 33 pl 7 figs 9-11 13-15











Fig 6


































Fig 6F-H

































































Fig 7

























































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References


















































































Fig 5J-K








Fig 5L










Doyle p 33 pl 7 figs 9-11 13-15











Fig 6


































Fig 6F-H

































































Fig 7

























































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References















































































Fig 6


































Fig 6F-H

































































Fig 7

























































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References















































































































Fig 6F-H

































































Fig 7

























































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References

















































































































Fig 7

























































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References






































































































































Fig 8







Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References




















































































Doyle 1990)















































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References






























































































al p 28 pl 2 fig 7














5 Discussion







Acknowledgements




References
















































































5 Discussion







Acknowledgements




References
















































































References














































































PINARD J.D., WEIS R., NEIGE P., MARIOTTI N. & DI CENCIO A. (2014) - Belemnites from the Upper...

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