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

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Kolumnentitel links: Jean-Daniel Pinard et al. Kolumnentitel rechts: Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous N. Jb. Geol. Paläont. Abh. 273/2 (2014), 155–177 Article E E Stuttgart, August 2014

N. Jb. Geol. Paläont. Abh. Article Stuttgart, August 2014

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, biostratigraphy 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 section 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 the Toarcian is also an important period of change in belemnite distribution, and concludes that a broad cor- Belemnites s.s. originated at the beginning of the Early relation between European areas and a notable ende- Jurassic (Doyle 1994; Riegraf 1996a; Weis & Del- mism for Siberia occurs. However, his study was at the sate 2006), though recent findings from Japan pro- generic level and focuses on the northern part of the vide arguments for a Triassic origin (Iba et al. 2012). Northwest European Province, and on Siberia. Finally, They display a successful diversification during the and even today, modern data for Toarcian belemnites Jurassic, with a peak in species richness for the Toar- are largely concentrated on the northern part of the cian. Detailed monographs have recently been pub- Northwest European Province (Germany, Great Brit- lished providing taxonomic and stratigraphic revised ain, Luxembourg, etc.) whereas little is known from data (e.g. Doyle 1990, 1992; Riegraf et al. 1984; the southern part of the Northwest European Pro- Sanders et al. 2013; Weis & Mariotti 2007; Weis et vince (Causses and Quercy basins, Provence Platform, al. 2012). However, these modern taxonomic revisions northern Spain, etc.), and from the Mediterranean are based on studies that are far from being homogene- Province (Italy, Austria, Southern Spain, North Africa, ous considering their palaeogeography. The Toarcian etc.). In all these areas, data are still dramatically low, (Early Jurassic), which concentrates most modern be- and what is known about belemnites is whether old lemnite taxonomic revisions, is exemplary. It is an im- and without any figuration or sparse (e.g. some isolat- portant period of taxonomic modification for belem- ed specimens figured, seeL issajous 1925; Sciau 1993, nites. Doyle’s synthesis paper (1994) points out that 2011; Combémorel in Rulleau et al. 1998). This lack

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).

of data is due to two factors (1) the low number of tion of belemnites to a standard chronostratigraphic belemnites study in these areas and (2) some outcrops frame based on ammonites. The reported belemnites (i.e. in Italy and Spain) are hardened carbonates lead- substantially increase the knowledge of Early Jurassic ing a very difficult extraction of well-preserved speci- faunas from the southern part of the Northwest Euro- mens. It is therefore impossible at the present day to pean Province, a step that could later lead to further portray the Jurassic belemnite diversification at the palaeogeographic explorations of the Jurassic radia- species level, mostly because we lack data from some tion. palaeogeographic areas, unless introducing strong palaeogeographic biases. 2. Geological and palaeogeographic In the present study we report a belemnite fauna settings of more than 700 specimens from the Tournadous section in the Causses Basin. This fauna constitutes We study here a single section (Tournadous) which ex- then a homogeneous and comprehensive record of poses Lower Jurassic marls located in Central-South the Late Pliensbachian and Toarcian belemnite fauna France (Fig. 1). This section has been studied by sev- in the southern part of the Northwest European Prov- eral authors, principally for its ammonoids content ince. These specimens have been collected in a com- (e.g. Monestier 1921, 1931; Guex 1972; Meister 1989; mon field study associating ammonites and belemnites Morard 2004) or to explore environmental changes scholars. It allows to assign the stratigraphic distribu- at the Upper Pliensbachian-Lower Toarcian bound- Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 157

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

ary (Mailliot et al. 2009). The studied portion in the proxies may be found in an extensive literature (e.g. present analysis covers the uppermost Pliensbachian Trümpy 1983; Graciansky et al. 1998; Mailliot et al. and continues up to the top of the Toarcian (see below). 2009; Van de Schootbrugge et al. 2013). Subsidence This section belongs to the Causses Basin, consid- is more or less pronounced within the basin leading ered as a small intracratonic basin that was part of an to variable thickness of the different formations and epicontinental sea. The basin was stuck between the even to hiatuses (see Trümpy 1983; Graciansky et al. “Massif Central” lands on the North and the “Mon- 1998). The studied section (Tournadous) is considered tagne Noire” lands on the South. Detailed analysis of as very proximal (Mailliot et al. 2009). the structure of the basin, sequence stratigraphic con- The studied interval covers three successive local text and stratigraphical correlations or geochemical formations. At the base of the section, the “Marnes de 158 Jean-Daniel Pinard et al.

Fig. 3. Stratigraphic repartition of collected ammonites and chronostratigraphic scheme of the section (Part 2). Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 159

Fig. 4. Stratigraphic repartition of the collected belemnites. 160 Jean-Daniel Pinard et al.

Fig. 5. Belemnites from Tournadous: Hastitidae. The specimens are all in natural size except where stated otherwise. A – Pleurobelus compressus (Stahl, 1824), BEL-256, rostrum in ventral view (A1), lateral view (A2) and dorsal view (A3). B – Pleurobelus compressus (Stahl, 1824), BEL-256, rostrum in ventral view (B1), lateral view (B2) and dorsal view (B3). C – 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 – Pleurobelus lagenaeformis (Hartmann in v. Zieten, 1832), BEL-260, rostrum in ventral view (E1), lateral view (E2) and dorsal view (E3). F – Pleurobelus lagenaeformis (Hartmann in v. Zieten, 1832), BEL-272, medial section of the rostrum (enlargement 2x). G – Pleurobelus lagenaeformis (Hartmann in v. Zieten, 1832), BEL-260, rostrum in ventral view (G1), lateral view (G2) and dorsal view (G3). H – Pleurobelus subirregularis (Lissajous, 1927), BEL-260, rostrum in ventral view (H1), lateral view (H2) and dorsal view (H3). I – Pleurobelus subirregularis (Lissajous, 1927), BEL-260, rostrum in ventral view (I1), lateral view (I2) and dorsal view (I3). J – Rhabdobelus donovani Riegraf in Riegraf et al., 1998, BEL-299, rostrum in ventral view (J1), lateral view (J2) and dorsal view (J3). K – Rhabdobelus donovani Riegraf in Riegraf et al., 1998, BEL-300, rostrum in ventral view (K1), lateral view (K2) and dorsal view (K3). L – Rhabdobelus aff. donovani Riegraf in Riegraf et al., 1998, BEL-293, rostrum in ventral view (L1), lateral view (L2) and dorsal view (L3).

Villeneuve” formation displays dark marls with a typi- within this formation are generally well-preserved. cal three nodular beds succession at its topmost part. Belemnites frequently bear their phragmocone and The last nodular bed has been considered to mark the ammonites are pyritized. Above is the well-known top of the Pliensbachian (Mailliot et al. 2009). Fossils “Schistes cartons” formation. It consists in finely lami- Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 161 nated shale sometimes containing silty material. Most which is thus probably present in the Tournadous sec- ammonites are compressed along laminations. Rare tion. All Toarcian chronozones are documented along pyritised nuclei may be found. Finally the “Marnes de the section, except for the Tenuicostatum Chronozone Fontaneilles” formation expands up to the Aalenian. (lowermost chronozone of Toarcian stage). This fits It consists mainly of marls with rare intercalated car- well previously published papers which claim that the bonate beds. Within the Tournadous section, a very entire Causses Basin lacks deposits from this lower- distinctive calcareous bed (bed number 100 here, see most Toarcian interval (Trümpy 1983; Guex et al. Fig. 2) occurs at nearly 20 meters above the base. Am- 2001; Mailliot et al. 2009). Within the section we monites from this formation are pyritised at the base note a distinctive condensed interval (bed 100, Figs. and calcareous and sometimes slightly compressed at 3-4) with the existence of fossil accumulation dated the top. Within this “Marnes de Fonteneilles” forma- from the Thouarsense Chronozone and from the base tion, gastropods and rare bivalves have been found. of the Dispansum Chronozone (Insigne Subchrono- Belemnites are frequent, well-preserved, mostly with zone). Interestingly, this accumulation at that age is their phragmocone. recorded by Graciansky et al. (1998: 617) in the much Based on ammonites palaeogeographic distribu- closed Quercy Basin, but not – as far as we know – in tions, Elmi et al. (1997) considered the Causses Basin other Causses Basin sections. This is probably due to to be southern part of the Northwest European Prov- the relatively proximal position of the Tournadous sec- ince, whereas for Page (2003) it was part of a Submedi- tion, contrary to the other more distal sections of the terranean Province for the beginning of the Toarcian basin. (Serpentinum and Bifrons chronozones) and part of the Northwest European Province after (from Variabi- 4. Systematic palaeontology lis to Aalensis chronozones). The systematics herein used is derived from Doyle (1994) and Riegraf (2000). Terminology follows Doyle & Kelly 3. Chronostratigraphy (1988). All studied belemnites are housed at the National Museum of Natural History, Luxembourg (MNHNL). Col- Ammonites have been precisely located along the sec- lected ammonites are housed at the University of Burgundy, tion (Figs. 2-3). From these distributions a chronos- France (UBGD, coll. Neige and Pinard). The belemnite ma- tratigraphic framework is proposed at the chronozone terial is composed of 710 specimens including 131 indeter- and subchronozone scales (Figs. 2-4). For that, we ex- minate rostra, 115 determinate only at the genus level and 464 at the species level. Detailed stratigraphic repartition of ommer plicitly refer to the landmark publications of D - the different species is given on Fig. 4. 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 Subclass Coleoidea Bather, 1888 where chronozones (and subchronozones) are defined Order Belemnitida Zittel, 1895 only at their base (see Page 2003). Question marks in Suborder Belemnitina Zittel, 1895 the present chronostratigraphic framework denote our Family Hastitidae Naef, 1922 impossibility to correctly date the referring beds due Genus PleurobelusNaef, 1922 to the absence of ammonites. The choice of a standard (i.e. chronostratigraphic) zonation is opportune Type species: Belemnites compressus Stahl, 1824. Upper because when established, this framework will serve Pliensbachian, Amaltheenton Formation, South-west Ger- as a reference to document temporal distribution of many. belemnite species. According to that approach, the studied part of Tournadous section expands from the Margaritatus Pleurobelus compressus (Stahl, 1824) Chronozone (Upper Pliensbachian) to the Aalensis Fig. 5A, B Chronozone (Upper Toarcian). We were not able to 1824 Belemnites compressus sp. n. – Stahl, p. 33. observe the Spinatum Chronozone (last Pliensbachian 1994 Pleurobelus compressus (Stahl, 1824). – Fischer, p. zone) (Fig. 2). However, Meister (1989) for different 13, pl. 2, fig. 3. sections in the Causses clearly recognizes it. Therefore 1998 Pleurobelus compressus (Stahl, 1824). – Schlegel- we probably missed index species or associated corre- milch, p. 54, pl. 4, fig. 3. lating fauna of the uppermost part of the Pliensbachian 162 Jean-Daniel Pinard et al.

Material: Thirty-six rostra from level 85, Upper Pliensbachi- Pleurobelus lagenaeformis (Hartmann in v. Zieten, an, Margaritatus Chronozone. MNHNL-BEL256, BEL257 1832) and BEL265. Fig. 5E-G Description: Small-sized, more or less slender, elongate 1832 Belemnites lagenaeformis Hartmann – v. Zieten, p. and laterally compressed rostrum with an obtuse apex. 33, pl. 25, fig. 1. Some individuals bear a reduced mucro. The outline is cy- 1848 Belemnites acuarius amalthei. – Quenstedt, p. 406 lindrical. The profile is asymmetrical, subhastate with a [pars], pl. 24, figs. 10, 13-14. more inflated venter in apical region. The transverse sec- 1998 Pleurobelus lagenaeformis (Hartmann). – Schle- tion is subquadrate to elliptical in the apical region with a gelmilch, p. 54, pl. 4, figs. 4-6. pronounced compression along the rostrum. There are no grooves. Lateral lines are more or less pronounced on the Material: Eighteen rostra from level 85, Upper Pliensbachi- anterior part of the rostrum. an, Margaritatus Chronozone. MNHNL-BEL260 to 262, BEL267 and BEL272. Remarks: See P. aff. compressus. Description: Small-sized cylindriconical orthorostrum Stratigraphic and geographic range: Upper Pliensbachi- with slightly obtuse apex. Some individuals bear a reduced an; England, Germany, France, Luxembourg, Switzerland, mucro. The outline and the profile are symmetrical and Slovakia. quite similar. The transverse section is rounded to elliptical and in the latter case slightly compressed. There are no grooves on the orthorostrum. The epirostrum is elongate Pleurobelus aff. compressus (Stahl, 1824) and tubular. Two dorso-lateral grooves are visible. The dis- Fig. 5C, D tinction between the epirostrum and the orthorostrum is well-pronounced. ?1848 Belemnites acuarius amalthei. – Quenstedt, p. 406 [pars], pl. 24, figs. 11-12. Remarks: The orthorostrum of P. lagenaeformis is similar to the rostrum of P. subirregularis. The only difference be- Material: Seven rostra from the level 85, Upper Pliensbachi- tween both is the presence of an epirostrum in P. lagenae- an, Margaritatus Chronozone. MNHNL-BEL258, BEL259, formis. Both species are described as distinct morphospe- BEL266 and BEL271. cies but could be considered as dimorphs from one biological species (cf. remark for P. aff. compressus). Description: Small-sized, more or less slender and elongate orthorostrum with an obtuse apex. Some individuals Stratigraphic and geographic range: Upper Pliensbachi- bear a reduced mucro. The outline is cylindrical. The pro- an; France, Germany. 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- Pleurobelus subirregularis (Lissajous, 1927) sion along the orthorostrum. There are no grooves on the Fig. 5H-I orthorostrum. Lateral lines are more or less pronounced on the anterior part of the orthorostrum. The epirostrum 1848 Belemnites acuarius amalthei. – Quenstedt, p. 406 is conical, its outline and profile are symmetrical with a [pars], pl. 24, fig. 9. slightly compressed section. Two dorso-lateral grooves and 1927 Dactyloteuthis subirregularis sp. n. – Lissajous, p. several striae are visible along the epirostrum. The distinc- 18, pl. 1, figs. 10-12. tion between the epirostrum and the orthorostrum is well- 1998 Pleurobelus subirregularis (Lissajous). – Schlegel- pronounced. milch, p. 55, pl. 4, fig. 7.

Remarks: The orthorostrum of P. aff. compressus is simi- Material: Sixteen rostra from level 85, Upper Pliensbachi- lar to the rostrum of P. compressus. The only difference lies an, Gibbosus Subchronozone. MNHNL-BEL263, BEL264, in the presence of an additional epirostrum in P. aff. com- BEL268 and BEL270. pressus. Both species are described as distinct morphospe- cies but could be considered as dimorphs from one biologi- Description: Small-sized cylindriconical rostrum with cal species, as stated by Doyle (1987) in the case of Cus- slightly obtuse apex. Some individuals bear a reduced piteuthis trivialis (Simpson, 1855) and C. tubularis (Young mucro. The outline and the profile are symmetrical and & Bird, 1922). P. aff. compressus differs from P. lagenae- similar. The transverse section is rounded to elliptical and formis by the morphology of their respective orthorostrum. in the latter case slightly compressed. There are no grooves.

Stratigraphic and geographic range: Upper Pliensbachi- Remarks: See P. lagenaeformis. an; France, Germany. Stratigraphic and geographic range: Upper Pliensbachi- an; France, Germany. Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 163

Genus Rhabdobelus Naef, 1922 Family Megateuthididae Sachs & Nalnjaeva, 1967 Genus Acrocoelites Lissajous, 1915 Type species: Belemnites exilis d’Orbigny, 1842. Toarcian iron ore of Saint Quentin-Fallavier (La Verpillière), Isère, Type species: Belemnites oxyconus Hehl in Zieten, 1831. France. Lower Toarcian, Posidonienschiefer Formation of Bad Boll, South-West Germany.

Rhabdobelus donovani Riegraf in Riegraf et al., 1998 Acrocoelites bobeti Lissajous, 1927 Fig. 5J-K Fig. 6A

1856 Belemnites parvus Hartmann. – Quenstedt, p. 286, 1927 Acrocoelites bobeti sp. n. – Lissajous, p. 19. pl. 41, fig. 21. 1971 Acrocoelites bobeti Lissajous. – Combémorel, p. 63, 1971 Belemnites parvus (Hartmann). – Schwegler, p. pl. 1, figs. 1-2. 115, text-figs. 119-120. 1990 Acrocoelites (Acrocoelites) bobeti. Lissajous. – 1998 Rhabdobelus donovani nom. nov. – Riegraf et al., p. Doyle, p. 33, pl. 7, figs. 9-11, 13-15. 85. Material: Seventeen rostra from the 106 and 107, Upper Material: One hundred and thirteen rostra from the level Toarcian, Pseudoradiosa Chronozone. MNHNL-BEL311 to 104 to 106, Upper Toarcian, Dispansum and Pseudoradiosa 316. chronozones. MNHNL-BEL299 to 309.

Description: Small-sized, slender and elongate rostrum Description: Medium-sized, slender and elongate rostrum with an acute apex. The outline is subhastate. The profile with acute apex. The outline and the profile are cylindrical is asymmetrical and hastate. The transverse section is sub- in the stem and conical in the apical region. However, the quadrate in the anterior part of the rostrum and also ventral- profile is slightly asymmetrical and moderately subhastate ly flattened in the alveolar region. A dorso-alveolar groove near the alveolar part. The transverse section is slightly extends until the anterior part of the rostrum solidum. This compressed and elliptical near the alveolar part and round- groove is more or less pronounced and sometimes replaced ed otherwise. Three apical grooves are visible, two dorso- by a flat area. Two well-developed lateral lines are visible laterals and one longer ventral. on the rostrum cavum and extend on the two thirds of the anterior part of the rostrum solidum. Stratigraphic and geographic range: Upper Toarcian; England, France, Germany. Stratigraphic and geographic range: Upper Toarcian; France, Germany. Acrocoelites brevisulcatus (Quenstedt, 1848) Fig. 6B Rhabdobelus aff. donovani Riegraf in Riegraf et al., 1998 1848 Belemnites acuarius brevisulcatus sp. n. – Quen- Fig. 5L stedt, p. 413, pl. 25, figs. 1, 3. 1969 Belemnites acuarius brevisulcatus Quenstedt. – Material: Twenty-nine rostra from the level 100 to 104, Up- Schwegler, p. 200, text-fig. 83. per Toarcian, Thouarsense and/or Dispansum chronozones. 1998 Acrocoelites (Odontobelus) brevisulcatus (Quen- MNHNL-BEL293 to 298 and BEL310. stedt). – Schlegelmilch, p. 62, pl. 8, figs. 4-5.

Description: Small sized, slender and elongate rostrum Material: Six rostra from the level 100, Upper Toarcian, with an acute apex. The outline is cylindriconical. The pro- Thouarsense and/or Dispansum chronozones. MNHNL- file is asymmetrical and slightly subhastate. The transverse BEL349 to 353. section is elliptical to slightly subquadrate. Two lateral lines are visible on the rostrum cavum and extend on the first Description: Small sized, short conical orthorostrum with third of the anterior part of the rostrum solidum. acute apex. The outline and the profile are similar and conical, with a slightly more inflated venter in the posterior Remarks: This species is distinguished from R. donovani part of the profile. The transverse section is compressed, by a short and conical rostrum, slightly subhastate in some elliptical to subquadrate. Three apical grooves are vis- individuals. The dorso-alveolar groove is always missing, ible, two dorso-laterals and a less pronounced ventral one. whereas well-developed in R. donovani. The epirostrum is elongate conical and bears on its entire length three grooves, the ventral one being more or less pro- Stratigraphic and geographic range: Upper Toarcian; nounced or absent in some individuals. Causses Basin, France. 164 Jean-Daniel Pinard et al.

Fig. 6. Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 165

Remarks: The orthorostrum of A. brevisulcatus is similar minstrensis (Fig. 6, C) (see also Riegraf et al. 1984: 155). A to the rostrum of A. pyramidalis. The only difference be- future revision will possibly elucidate this hypothesis. tween both is the presence of an epirostrum in A. brevisulcatus. Both species could be considered as two morpho- Stratigraphic and geographic range: Lower Toarcian; types of the same biological species. Bulgaria, England, France, Germany, Luxembourg.

Stratigraphic and geographic range: Upper Toarcian; Germany, France. Acrocoelites levidensis (Simpson, 1855) Fig. 6D

1855 Belemnites levidensis sp. n. – Simpson, p. 20. Acrocoelites ilminstrensis (Phillips, 1867) 1990 Acrocoelites (Odontobelus) levidensis (Simpson). – Fig. 6C Doyle, p. 45, pl. 14, fig. 8, pl. 15: 4-7. 2003 Belemnites levidensis Simpson [Acrocoelites (Od.) 1848 Belemnites tripartitus paxillosus. – Quenstedt, p. levidensis (Simpson)]. – Doyle, p. 175, fig. 15e-f. 420, pl. 26, figs. 25-26. 2003 Belemnites alveolatus Simpson [Acrocoelites (Od.) 1867 Belemnites ilminstrensis sp. n. – Phillips, p. 64, pl. levidensis (Simpson)]. – Doyle, p. 175, fig. 12g-h. 12, fig. 30. 1984 Acrocoelites (Acrocoelites) ilminstrensis (Hehl). – Material: Nine rostra from the level 92 to 96, Lower Toar- Riegraf et al., p. 155, pl.10, figs. 2-3, 5, 9. cian, Bifrons Chronozone. MNHNL-BEL340 to 346. 1990 Acrocoelites (Toarcibelus) ilminsterensis [sic] (Phillips). – Doyle, p. 39, pl. 1, figs. 1-7. Description: Medium- to large-sized and robust rostrum with acute apex. The outline is cylindriconical. The profile Material: Three rostra from the level 92, Lower Toarcian, is conical and asymmetrical with a slightly inflated ventral Bifrons Chronozone. MNHNL-BEL333 to 335. part. The transverse section is compressed and elliptical to slightly subquadrate. Three apical grooves are visible, two Description: Medium to large sized, more or less slender dorso-laterals and one ventral, with the latter more pro- and elongate rostrum with acute apex. The outline is cy- nounced and longer. lindriconical. The profile is conical to cylindriconical and symmetrical. The transverse section elliptical and slightly Stratigraphic and geographic range: Toarcian; England, compressed. Three apical grooves are visible, two dorso- France, Germany, Luxembourg. 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. Acrocoelites oxyconus (Hehl in v. Zieten, 1831) Fig. 6E Remarks: A. ilminstrensis shows a striking similarity with A. oxyconus (Hehl in v. Zieten, 1831). All the transitional 1831 Belemnites oxyconus Hehl. – v. Zieten, p. 27, pl. 21, morphologies are visible in our material and also in the fig- fig. 5. ured specimens in literature (Doyle 1990; Riegraf et al. 1848 Belemnites tripartitus oxyconus sp. n. – Quenstedt, 1984). In our opinion, these species are only based on the p. 419, pl. 26, fig.19. separation of two extreme morphotypes: A. oxyconus (Fig. 1984 Acrocoelites (Acrocoelites) oxyconus (Hehl). – Rie- 6E) being slightly more compressed and conical than A. il- 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 – Acrocoelites bobeti Lissajous, 1927, BEL-311, rostrum in ventral view (A1), lateral view (A2). B – Acrocoelites brevisulcatus (Quenstedt, 1848), BEL-349, rostrum in ventral view (B1), lateral view (B2). C – Acrocoelites ilminstrensis (Phil- lips, 1867), BEL-333, rostrum in ventral view (C1), lateral view (C2). D – Acrocoelites levidensis (Simpson, 1855), BEL-346, rostrum in ventral view (D1), lateral view (D2). E – Acrocoelites oxyconus (Hehl in v. Zieten, 1831), BEL-254, rostrum in ventral view (E1), lateral view (E2). F – Acrocoelites pyramidalis (Münster in v. Zieten, 1831), BEL-239, rostrum in ventral view (F1), lateral view (F2). G – Acrocoelites pyramidalis (Münster in v. Zieten, 1831), BEL-240, rostrum in ventral view (G1), lateral view (G2). H – Acrocoelites pyramidalis (Münster in v. Zieten, 1831), BEL-241, rostrum in ventral view (H1), lateral view (H2). I – Acrocoelites conoideus (Oppel, 1856), BEL-326, rostrum in ventral view (I1), lateral view (I2). J – Acrocoelites cf. riegrafi Doyle, 1992, BEL-338, rostrum in ventral view (J1), lateral view (J2). K – Acrocoelites rostriformis (Theodori in Bronn, 1837), BEL-242, rostrum in ventral view (K1), lateral view (K2). L – Acrocoelites rostriformis (Theodori in Bronn, 1837), BEL-243, rostrum in ventral view (L1), lateral view (L2). 166 Jean-Daniel Pinard et al.

1990 Acrocoelites (Acrocoelites) oxyconus (Zieten). – Description: Large and robust rostrum with moderately Doyle, p. 29, pl. 6, figs.1-4. acute apex. The outline is cylindriconical. The profile is cylindriconical and asymmetrical with a slightly inflated Material: Six rostra from the level 92 to 95, Lower Toar- ventral part. The transverse section is rounded to slightly cian, Bifrons Chronozone. MNHNL-BEL254, BEL255 and elliptical. Two dorso-lateral and one ventral apical grooves BEL142. are equally developed.

Description: Medium- to large-sized, more or less slender Stratigraphic and geographic range: Upper Toarcian to and elongate rostrum with acute apex. The outline is cy- Lower Aalenian; Bulgaria, Caucasus, France, Germany, lindriconical. The profile is conical to moderately cylindri- Luxembourg, Slovakia, and possibly Romania. conical and symmetrical. The transverse section is elliptical and compressed. Three apical grooves are visible, two dorso-laterals and a longer and more incised ventral one. There Acrocoelites cf. riegrafi Doyle, 1992 are no true lateral lines, only lateral flattenings are visible. Fig. 6J Remarks: See A. ilminstrensis. cf. 1969 Belemnites longiconus sp. n. – Schwegler, p. 187, Stratigraphic and geographic range: Lower Toarcian; text-fig. 73. England, France, Germany, Slovakia. cf. 1990 Acrocoelites (Acrocoelites) longiconus (Schweg- ler). – Doyle, p. 34, pl. 7, figs. 4-6, 17. cf. 1992 Acrocoelites (Acrocoelites) riegrafi nom. nov. – Doyle, p. 75. [pro A. (A.) longiconus (Schwegler)] Acrocoelites pyramidalis (Münster in v. Zieten, cf. 1998 Acrocoelites (Acrocoelites) riegrafi Doyle. – 1831) Schlegelmilch, p. 59, pl. 5, fig. 10. Fig. 6F-H Material:One rostrum from the Lower Toarcian, Bifrons 1831 Belemnites pyramidalis Münster – v. Zieten, p. 31, Chronozone. MNHNL-BEL338. pl. 24, fig. 5. 1998 Acrocoelites (Odontobelus.) pyramidalis (Mün- Description:Small-sized cylindriconical orthorostrum. The ster). – Schlegelmilch, p. 62, pl. 8, figs.1-2. outline and the profile are similar and cylindriconical. The 1998 Acrocoelites (Odontobelus) pyramidalis (Zieten). – transverse section is compressed elliptical. The epirostrum Combémorel in Rulleau et al., p. 21, pl. 1, fig. 8. is short, conical and bears two well-developed dorso-lateral grooves and one weakly developed ventral groove. Material: One hundred and nineteen rostra from the level 95 to 106, Lower to Upper Toarcian, Bifrons to Pseudora- Remarks: Our sample shows a strong resemblance with the diosa chronozones. MNHNL-BEL239 to 241, BEL244 to figured type specimen (Schlegelmilch 1998, pl. 5, fig. 10) 250 and BEL330. but differs by its reduced ventral groove.

Description: Small-sized, short conical rostrum with acute Stratigraphic and geographic range: Lower Toarcian; apex. The outline and the profile are similar and conical, Causses Basin, France (this report). 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- Acrocoelites rostriformis (Theodori in Bronn, 1837) laterals and a less pronounced ventral one. Fig. 6K-L Stratigraphic and geographic range: Lower to Upper Toarcian; England, France, Germany, Luxembourg, Slova- 1837 Belemnites rostriformis Theodori. – Bronn, p. 412. kia, possibly Caucasus. 1848 Belemnites rostriformis Theodori. – Quenstedt, p. 425, pl. 27, figs. 19-20. 1998 Acrocoelites (Odontobelus) curtus (d’Orbigny). – Schlegelmilch, p. 62, pl. 5, fig. 14, pl. 8, fig. 3. Acrocoelites conoideus (Oppel, 1856) 1998 Acrocoelites curtus (d’Orbigny) – Combémorel in Fig. 6I Rulleau et al., p. 29, pl. 2, figs. 8-10.

1848 Belemnites compressus conicus sp. n. – Quenstedt, Material: Nine rostra from the level 100 to 106, Upper p. 423, pl. 27, fig. 4. Toarcian, Thouarsense and/or Dispansum to Pseudoradiosa 1856 Belemnites conoideus sp. n. – Oppel, p. 483. chronozones. MNHNL-BEL242, BEL243, BEL251 to 253 1998 Acrocoelites (Acrocoelites) conoideus (Oppel) – and BEL347. Schlegelmilch, p. 60, pl. 6, figs.7-8. Description: Small-sized, short conical rostrum with an Material: One rostrum from the level 111, Upper Toarcien, acute apex. The outline is conical. The profile is conical Mactra Subchronozone. MNHNL-BEL326. and asymmetrical with an inflated ventral part. The trans- Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 167 verse section is rounded to elliptical in the anterior part and Stratigraphic and geographic range: Lower Toarcian; subpyriform in the apical region. Three apical grooves are England, France, Germany. visible, two well-developed dorso-lateral grooves and a ventral one, reduced in some individuals. Acrocoelites tripartitus (v. Schlotheim, 1820) Stratigraphic and geographic range: Upper Toarcian to Fig. 7C lowermost Aalenian; France, Germany, Luxembourg. 1820 Belemnites tripartitus sp. n. – v. Schlotheim, p. 48. [pars] Acrocoelites inaequistriatus (Simpson, 1855) 1942 Mesoteuthis banzensis sp. n. – Kolb, p. 163, pl. 11, Fig. 7A figs. 1-2, 7. 1996b Acrocoelites (Acrocoelites) tripartitus (v. Schlothe- 1855 Belemnites inaequistriatus sp. n. – Simpson, p. 24. im). – Riegraf, p. 27, fig. 24c. 1990 Acrocoelites (Toarcibelus) inaequistriatus (Simp- 2000 Acrocoelites tripartitus (v. Schlotheim). – Riegraf, son). – Doyle, p. 41, pl. 13, figs. 1-5, pl. 14, figs. 1-3, p. 287, figs. 3-4, 18. 7. 1998 Acrocoelites (Toarcibelus) inaequistriatus (Simp- Material: Twelve rostra from the level 100 to 109, Upper son). – Rulleau et al., p. 24, pl. 2, fig. 1. Toarcian, Variabilis to Aalensis chronozones. MNHNL- 2003 Belemnites inaequistriatus Simpson [Acrocoelites BEL322 to 325. (Praepachyteuthis) inaequistriatus (Simpson)]. – Doyle, p. 173, fig. 13a-b. Description: Medium-sized, conical to cylindriconical rostrum with an acute apex. The outline is conical. The profile Material: Two rostra from the level 100, Upper Toarcian, is conical to cylindriconical and asymmetric with a moder- Thouarsense and/or Dispansum chronozones. MNHNL- ately inflated venter. The transverse section is moderately BEL336. elliptical compressed. Three apical grooves, two dorso-laterals and one longer and more incised ventral, are visible. Description: Medium- to large-sized, conical rostrum with epirostrum. The outline and the profile of the entire rostrum Remarks: The present species can be distinguished from are similar and conical. The transverse section is moderate- A. conoideus and A. quenstedti (Oppel, 1856) by its more ly elliptical compressed to rounded. Irregular longitudinal elongate and conical apical region. striae and three well-developed apical grooves (two dorso- lateral and one ventral) are visible on the epirostrum. Stratigraphic and geographic range: Upper Toarcian; France, Germany, Luxembourg possibly Ukraine and Cau- Stratigraphic and geographic range: Lower to Upper casus. Toarcian; England, France, Luxembourg.

Acrocoelites vulgaris (Young & Bird, 1822) Acrocoelites subtenuis (Simpson, 1855) Fig. 7D Fig. 7B 1822 Belemnites vulgaris sp. n. – Young & Bird, p. 258, 1855 Belemnites subtenuis sp. n. – Simpson, p. 26. pl. 14, fig. 1. 1990 Acrocoelites (Acrocoelites) subtenuis (Simpson). – 1867 Belemnites vulgaris Young & Bird. – Phillips, p. 73, Doyle, p. 30, pl. 6, figs. 5, 7-14, pl. 7, figs. 1-3, text- pl. 16, figs. 40-41. fig. 15. 1992 Acrocoelites (Odontobelus) vulgaris (Young & 2003 Belemnites subtenuis Simpson [Acrocoelites (Acro- Bird). – Doyle, p. 44, pl. 11, figs. 6-8, pl. 13, figs. coelites) subtenuis (Simpson)]. – Doyle, p. 169, fig. 6-7, pl. 14, figs. 4-6, pl. 15, figs. 2-3. 9c-f, k-l. Material: Two rostra from the levels 95 and 98, Lower Material: Six rostra from the level 91-92, Lower Toarcian, to Upper Toarcian, Bifrons to Variabilis chronozones. Serpentinum to Bifrons chronozones. MNHNL-BEL327 to MNHNL-BEL331, BEL 32 and BEL339. 329. Description: Large-sized, robust and cylindriconical ros- Description: Medium-sized, slender and elongate cylindri- trum with a slightly recurved acute apex. The outline is cy- conical rostrum. The outline is cylindriconical and slightly lindriconical. The profile is symmetrical and conical. The subhastate. The profile is symmetrical and cylindriconical. transverse section is moderately compressed and elliptical. The transverse section is slightly elliptical compressed near Three apical grooves, two dorso-laterals and one longer and the alveolar part, rounded in the medium part and slightly more incised ventral, are visible. depressed subpyriform in the apical region. The venter of the apical region is moderately flattened. Three apical grooves Stratigraphic and geographic range: Toarcian; England, are visible, two short dorso-laterals and one longer ventral. France, Germany, Luxembourg. 168 Jean-Daniel Pinard et al.

Fig. 7. Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 169

Genus Cuspiteuthis Abel, 1916 Genus Dactyloteuthis Bayle, 1878

Type species: Belemnites acuarius v. Schlotheim, 1820. Type species: Belemnites irregularis v. Schlotheim, 1813. Lower Toarcian (Falciferum Subzone), Bituminous Shales Upper Toarcian (Variabilis Zone), Jurensismergel Forma- Formation, Saltwick Bay, North Yorkshire. tion, Upper Franconia (South Germany) (fide Riegraf 2000).

Cuspiteuthis acuaria (v. Schlotheim, 1820) Fig. 8A Dactyloteuthis incurvata (v. Zieten, 1831) Fig. 7E 1820 Belemnites acuarius sp. n. – v. Schlotheim, p. 46. [pars]. 1831 Belemnites incurvatus sp. n. – v. Zieten, p. 29, pl. 1848 Belemnites acuarius gigas sp. n. – Quenstedt, p. 22, fig. 7a-e. 412, pl. 25, figs. 6-7. 1984 Dactyloteuthis (Dactyloteuthis) incurvata (v. Zie- 1984 Youngibelus gigas (Quenstedt). – Riegraf et al., p. ten). – Riegraf et al., p. 161, pl. 12:, fig.10, text-fig. 158, pl. 11, figs. 9-11. 48r. 1998 Youngibelus ohmdenensis nom. nov. – Schlegel- 1998 Belemnites pygmeus v. Zieten [Dactyloteuthis]. – milch, p. 63, pl. 8, figs. 10-11. Riegraf et al., p. 217. 2000 Cuspiteuthis acuaria (v. Schlotheim). – Riegraf, p. 1998 Dactyloteuthis incurvata (v. Zieten). – Schlegel- 292, figs. 9-10, 13-14. milch, p. 66, pl. 9, figs. 12-13.

Material: Five rostra from the level 95, Lower Toarcian, Bi- Material: Two rostra from the Lower Toarcian, Bifrons frons Chronozone. MNHNL-BEL92, BEL291 and BEL292. Chronozone. MNHNL-BEL280 and BEL281.

Description: Medium-sized, conical orthorostrum with a Description: Small-sized and robust rostrum with an acute more or less acute apex and a well-developed epirostrum. apex. The outline is cylindriconical to slightly subhastate. The outline and the profile of the orthorostrum are conical. The profile is symmetrical and cylindriconical. The trans- The transverse section is rounded to elliptical and slightly verse section is well-compressed, rounded subquadrate and compressed. No grooves are visible on the orthorostrum. slightly pyriform in the apical region. Three apical grooves The epirostrum is very elongate and conical to tubular in are visible, the ventral one is well-developed contrary to the posterior part. the dorso-laterals which are more or less incised and do not reach the tip of the apex. Remarks: C. acuaria shows a striking similarity with C. tubularis (Young & Bird, 1822). All the transitional mor- Remarks: Riegraf et al. (1998: 78) stated that Belemnites phologies are visible in our material and also in the figured incurvatus v. Zieten, 1831 is a primary homonym of Be- specimens in literature (Doyle, 1992; Riegraf et al. 1984). lemnites incurvatus Raspail, 1829 and replace it by Belem- In our opinion, these species are certainly based on the nites pygmeus v. Zieten, 1831, whose holotype has been re- separation of two extreme morphotypes: C. acuaria being studied by Riegraf (in Riegraf et al. 1998: 78). However, slightly more robust and conical than C. tubularis. A future Belemnites pygmeus is possibly a nomen nudum and/or no- revision will possibly elucidate this hypothesis. men oblitum (personal communication W. Riegraf 2003) and the better-known name Belemnites incurvatus is herein Stratigraphic and geographic range: Lower to lowermost employed, in anticipation of further clarification. Upper Toarcian; France, Germany, Luxembourg. 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 – Acrocoelites inaequistriatus (Simpson, 1855), BEL-336, rostrum in ventral view (A1), lateral view (A2). B – Acrocoe- lites subtenuis (Simpson, 1855), BEL-327, rostrum in ventral view (B1), lateral view (B2). C – Acrocoelites tripartitus (v. Schlotheim, 1820), BEL-322, rostrum in ventral view (C1), lateral view (C2). D – Acrocoelites vulgaris (Young & Bird, 1822), BEL-339, rostrum in ventral view (D1), lateral view (D2). E – Dactyloteuthis incurvata (v. Zieten, 1831), BEL-280, rostrum in ventral view (E1), lateral view (E2). F – Dactyloteuthis irregularis (v. Schlotheim, 1813), BEL-282, rostrum in ventral view (F1), lateral view (F2). G – Dactyloteuthis cf. semistriata (Münster, 1830), BEL-286, rostrum in ventral view (G1), lateral view (G2). H – Dactyloteuthis similis (v. Seebach, 1864), BEL-288, rostrum in ventral view (H1), lateral view (H2). I – Dactyloteuthis wrighti (Oppel, 1856), BEL-348, rostrum in ventral view (I1), lateral view (I2). 170 Jean-Daniel Pinard et al.

Dactyloteuthis irregularis (v. Schlotheim, 1813) pressed and elliptical. No apical grooves are visible on our Fig. 7F specimen and the epirostrum is missing, just the scar is visible. 1813 Belemnites irregularis sp. n. – v. Schlotheim, p. 70, pl. 3, fig. 2. Remarks: The preservation of the unique specimen does 1998 Dactyloteuthis irregularis (v. Schlotheim). – Schle- not allow a definitive determination. The characteristic gelmilch, p. 65, pl. 9, figs. 1-4. epirostrum is not preserved, only a scar indicates its pres- 2000 Dactyloteuthis irregularis (v. Schlotheim). – Rie- ence. graf, p. 293, figs. 1-2, 17. Stratigraphic and geographic range: Upper Toarcian; Material: Ten rostra from the level 100, Upper Toarcian, France (this report). Thouarsense Chronozone and/or Insigne Subchronozone. MNHNL-BEL282 to 285. Dactyloteuthis similis (v. Seebach, 1864) Description: Medium-sized, robust and digit-like rostrum Fig. 7H with very obtuse apex. The outline and the profile are cylindrical to cylindriconical. The transverse section is strongly 1864 Belemnites similis sp. n. – v. Seebach, p. 158, pl.7, compressed and elliptical. The presence and the size of the fig. 6. apical ventral groove are variable. 1998 Dactyloteuthis irregularis (Schlotheim). – Rulleau et al., p. 25, pl. 2, figs. 2-3. Stratigraphic and geographic range: Upper Toarcian; 1998 Dactyloteuthis similis (v. Seebach). – Schlegel- Bulgaria, England, France, Germany, Luxembourg, Poland, milch, p. 65, pl. 9, figs. 5-6. Slovakia. Material: Five rostra from the level 100, Upper Toarcian, Thouarsense and/or Dispansum chronozones. MNHNL- Dactyloteuthis cf. semistriata (v. Münster, 1830) BEL287 to 289. Fig. 7G Description: Medium-sized, slightly elongate and digit-like rostrum with very obtuse apex. The outline and the profile cf. 1830 Belemnites semistriatus sp. n. – Münster, p. 15, pl. 2, fig. 4. are cylindrical to cylindriconical. The transverse section is distinctly compressed and elliptical. There is no apical cf. 1848 Belemnites acuarius ventricosus sp. n. – Quen- groove. stedt, p. 411, pl. 25, fig. 8. cf. 1984 Dactyloteuthis (Cuspiteuthis) semistriata (Mün- Stratigraphic and geographic range: Upper Toarcian; ster). – Riegraf et al., p. 163, pl. 12, fig. 4, 12, France, Germany, Luxembourg. text-fig. 48u. cf. 1994 Dactyloteuthis irregularis (Schlotheim) [Belem- nites acuarius]. – Fischer, p. 9, pl. 1, figs. 4-5. Dactyloteuthis wrighti (Oppel, 1856) Material: One adult rostrum from the level 100, Upper Fig. 7I Toarcian, Thouarsense and/or Dispansum chronozones. MNHNL-BEL286. 1848 Belemnites digitalis tripartitus sp. n. – Quenstedt, p. 418, pl. 26, figs. 14, 31. Description: Small-sized and digit-like orthorostrum with 1856 Belemnites wrighti sp. n. – Oppel, p. 240. very obtuse apex. The outline and the profile are cylindrical 1990 Acrocoelites (Odontobelus) wrighti (Oppel, 1856). – to cylindriconical. The transverse section is strongly com- 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 – Cuspiteuthis acuaria (v. Schlotheim, 1820), BEL-92, rostrum in ventral view (A1), lateral view (A2). B – Parapassaloteuthis cf. polita (Simpson, 1866), BEL-278, rostrum in ventral view (B1), lateral view (B2). C – Passaloteuthis bruguieriana (d’Orbigny, 1842), BEL-275, rostrum in ventral view (C1), lateral view (C2). D – Passaloteuthis laevigata (v. Zieten, 1831), BEL-273, rostrum in ventral view (D1), lateral view (D2). E – Passalo- teuthis milleri (Phillips, 1867), BEL-277, rostrum in ventral view (E1), lateral view (E2). F – Pseudohastites sp., BEL-277, rostrum in ventral view (F1), lateral view (F2). G – Salpingoteuthis dorsetensis (Oppel, 1856), BEL-357, rostrum in ventral view (G1), lateral view (G2) and dorsal view (G3). H – Salpingoteuthis trisulcata (de Blainville, 1827), BEL-317, rostrum in ventral view (H1), lateral view (H2) and dorsal view (H3). Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 171

Fig. 8. 172 Jean-Daniel Pinard et al.

1994 Dactyloteuthis incurvata (Zieten) [Belemnites no- Passaloteuthis bruguieriana (d’Orbigny, 1842) dotianus (d’Orbigny)] – Fischer, p. 13, pl. 2, figs. Fig. 8C 5-6. 1842 Belemnites bruguierianus sp. n. – d’Orbigny, p. 84. Material: Two rostra from the Lower Toarcian, Bifrons 1990 Passaloteuthis bisulcata (Blainville) [variety A]. – Chronozone. MNHNL-BEL348. Doyle, p. 19, pl. 1, figs. 1-2, pl. 2, fig. 1. 1990 Passaloteuthis bisulcata (Blainville) [variety B]. – Description: Medium-sized and elongate rostrum with Doyle, p. 19, pl. 2, figs. 2-3. acute apex. The outline and the profile are cylindriconical. 1998 Passaloteuthis bisulcata (Blainville). – Schlegel- The transverse section is compressed and elliptical. There is milch, p. 51, pl. 2, figs. 6-7. one short, more or less incised, ventral groove. Material: Two rostra from the level 85, Upper Pliensbachi- Stratigraphic and geographic range: Toarcian; England, an, Margaritatus Chronozone. MNHNL-BEL275 and France, Germany, Greenland, Luxembourg. BEL276.

Description: Large-sized, robust cylindriconical rostrum Family Passaloteuthididae Naef, 1922 with a more or less obtuse apex. The outline and the profile Genus Parapassaloteuthis Riegraf, 1980 (emend. are cylindriconical. The transverse section is rounded. Two Doyle, 1990) weakly developed dorso-lateral apical grooves are present.

Type species: Belemnites zieteni Werner, 1912 [= Belem- Stratigraphic and geographic range: Upper Pliensbachian nites zieteni Mayer-Eymar, 1884; fide Riegraf et al. 1998]. to lowermost Toarcian; Europe (Bulgaria, England, France, Upper Pliensbachian, Amaltheenton Formation, South-west Germany, Italy, Luxembourg, Portugal, Slovakia, Spain), Germany. northern Africa (Algeria, Morocco), Western Asia (Turkey).

Parapassaloteuthis cf. polita (Simpson, 1866) Passaloteuthis laevigata (v. Zieten, 1831) Fig. 8B Fig. 8D cf. 1866 Belemnites politus sp. n. – Simpson, p. 216. 1831 Belemnites laevigatus sp. n. – v. Zieten, p. 28, pl. 21, cf. 1990 Parapassaloteuthis polita (Simpson). – Doyle, p. fig. 12. 27, pl. 5, figs. 3-9. 1990 Passaloteuthis bisulcata (Blainville) [variety C]. – cf. 2003 Belemnites politus Simpson [Parapassaloteuthis Doyle, p. 19, pl. 3, figs. 1-3. polita (Simpson)]. – Doyle, p. 167, fig. 6m-n. 1998 Passaloteuthis laevigata (Zieten) – Schlegelmilch, p. 51, pl. 2, fig. 8. Material: One rostrum from the level 85, Upper Pliensba- chian, Margaritatus Chronozone. MNHNL-BEL278. Material: Six rostra from the level 85, Upper Pliensbachian, Margaritatus Chronozone. MNHNL-BEL273 and BEL274. Description: Small-sized, robust rostrum with a slightly mucronate apex. The outline is cylindriconical. The pro- Description: Large-sized, elongate cylindriconical rostrum file is cylindriconical and asymmetrical with more inflated with a moderately acute apex. The outline and the profile venter. The transverse section is slightly compressed and are cylindriconical to slightly conical. The transverse sec- rounded subquadrate. Two dorso-lateral apical grooves are tion is rounded subquadrate. Two well-developed dorso- weakly developed. lateral apical grooves are present.

Remarks: Our specimen shares morphological charac- Stratigraphic and geographic range: Upper Pliensbachi- ters of Pa. brevis (de Blainville, 1827) from the Lower an to lowermost Toarcian; England, France, Germany, Lux- Pliensbachian and Pa. robusta (Simpson, 1855) from the embourg. Lower Toarcian. It resembles most closely some Pa. polita in Doyle (1990, pl. 5, figs. 6-7) but lacks the typical subhastate outline, as given in the diagnosis (Doyle 1990: 27). Passaloteuthis milleri (Phillips, 1867) Stratigraphic and geographic range: Upper Pliensbach- Fig. 8E ian; France (this report). 1867 Belemnites milleri sp. n. – Phillips, p. 54, pl. 8, fig. 19. 1992 Passaloteuthis milleri (Phillips). – Doyle, p. 22, pl. Genus Passaloteuthis Lissajous, 1915 2, figs. 5-9. 1998 Passaloteuthis milleri (Phillips). – Schlegelmilch, Type species: Belemnites bruguierianus d rbigny ’O , 1842. p. 52, pl. 3, fig. 4, pl. 7, fig. 6. Pliensbachian, France. Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 173

Material: Two rostra from the level 90, Upper Pliensbach- Material: Two rostra from the level 106, Upper Toar- ian. MNHNL-BEL277. cian, Pseudoradiosa Chronozone. MNHNL-BEL357 and BEL358. Description: Small-sized cylindrical rostrum with an obtuse apex. The outline is cylindrical and the profile cylindri- Description: Short reduced conical orthorostrum with conical. The transverse section is rounded to subpyriform acute apex. The outline is conical. The profile is asymmet- near the apex. No apical grooves are visible. rical and conical. The transverse section is rounded. The epirostrum is strongly developed (90% of the total rostrum) Stratigraphic and geographic range: Upper Pliensbach- and conical to short tubular. A broad apical ventral groove, ian to lowermost Toarcian; Bulgaria, England, France, Ger- several apical striae and lateral lines, fading out near the many, Luxembourg. apex, are visible.

Stratigraphic and geographic range: Upper Toarcian to Aalenian; Bulgaria, Caucasus, England, France, Germany. Genus Pseudohastites Naef, 1922

Type species: Belemnites scabrosus Simpson, 1866. Lower Pliensbachian (Jamesoni Zone), Robin Hood’s Bay, North Salpingoteuthis trisulcata (de Blainville, 1827) Yorkshire. Fig. 8H

Pseudohastites sp. 1827 Belemnites trisulcatus sp. n. – de Blainville, p. 83, Fig. 8F pl. 5, fig. 13. 1848 Belemnites acuarius tricanaliculatus sp. n. – Quen- Material: One adult rostrum from the level 85, Up- stedt, p. 414, pl. 25, figs. 13-14. per Pliensbachian, Margaritatus Chronozone. MNHNL- 1992 Salpingoteuthis trisulcata (Blainville). – Doyle, p. BEL337. 71, pl. 28, figs. 2-5, 10. 1994 Salpingoteuthis trisulcata (Blainville). – Fischer, Description: Large-sized, slender and elongate orthoros- p. 13, pl. 2, figs. 12-13. trum with a moderately acute apex. The outline and the profile are similar and subhastate. The transverse section is Material: Twelve rostra from the level 109 and 110, Upper rounded subquadrate in the anterior part of the rostrum to Toarcian, Aalensis Chronozone. MNHNL-BEL317 to 321. rounded in the posterior part. A strong dorso-alveolar flattening is present. The epirostrum is conical to subtubular Description: Short reduced conical orthorostrum with and bears two shallow dorso-lateral grooves. acute apex. The profile is conical. The transverse section is rounded. The epirostrum is strongly developed (90% of Remarks: The specimen fits well the diagnosis of Pseu- the total rostrum) and very elongate tubular. The dorso- lateral grooves, paired in the anterior part, are always well- dohastites (Doyle 1990: 23) but cannot be referred to any known species for its peculiar characters: the strong dorso- developed. The incision of the dorsal and ventral grooves, alveolar flattening and a true epirostral development. extending to the alveolar region, is highly variable. Several irregular striae are present. Stratigraphic and geographic range: Upper Pliensbachi- an; France (this report).

Family Salpingoteuthididae Doyle, 1992 Genus Salpingoteuthis Lissajous, 1915

Type species: Belemnites trisulcatus de Blainville, 1827. Toarcian, Calvados, Northwest France.

Salpingoteuthis dorsetensis (Oppel, 1856) Fig. 8G Fig. 9. Specific richness throughout the studied period. 1856 Belemnites dorsetensis sp. n. – Oppel, p. 362. The condensate level is represented by a single bar for the 1992 Salpingotheuthis dorsetensis (Oppel). – Doyle, p. Thouarsense Chronozone and the beginning of the Dispan- 72, pl. 27, figs. 1-2. sum Chronozone, by the fact of the melting of both faunas 1998 Salpingotheuthis dorsetensis (Oppel). – Rulleau et (Ma: Margaritatus, Sp: Spinatum, Te: Tenuicostatum, Ser- al., p. 28, pl. 2, fig. 7. pentinum, Bi: Bifrons, Va: Variabilis, Th: Thouarsense, Di: Dispansum, Ps: Pseudoradiosa, Aa: Aalensis). 174 Jean-Daniel Pinard et al.

Stratigraphic and geographic range: Upper Toarcian to rather abundant in the northern part of the Northwest Aalenian; England, France, Germany, Slovakia. European Province (England and Southern Germany). A similar pattern is observed for Southern France in 5. Discussion the Aalenian-Bajocian (see Mariotti et al. 2012). In the same way, there are relatively few Dactyloteuthis The analysis of the belemnite fauna reveals that specif- in Tournadous. On the contrary there is a predomi- ic richness is not constant over the studied period (Fig. nance of the genera Rhabdobelus and Salpingoteuthis 9). Due to lack of data, the diversity variations during in the late Toarcian, a pattern which is quite differ- the Late Pliensbachian cannot be described precisely. ent from the northern part of the Northwest European However, it is possible to observe an important spe- Province (Doyle 1992; Weis 1999). These apparent cies richness decrease between the Margaritatus and absences/predominances of the mentioned genera hint Serpentinum chronozones. From the Serpentinum to at an onsetting provincialism during the Toarcian, Bifrons chronozones, species richness rapidly rises, with “southern, or Tethyan, belemnite faunas” as sug- reaching a maximum during the Bifrons Chronozone. gested by Doyle (1987, 1994) and subsequently for the Subsequently, the species richness strongly decreases Aalenian by Mariotti et al. (2012). (in Variabilis Chronozone) and then recovers, reach- Based on the study of a single outcrop, it is impos- ing a peak during Thouarsense Chronozone (or the sible, at the present day, to make a detailed compari- beginning of Dispansum Chronozone) with similar son with the well-known faunas of the northern part levels than previously seen in the Bifrons Chrono- of the Northwest European Province (such as done zone. It is, however, not possible to be more precise for the same geological ages by Dera et al. 2011 for on the position of this peak due to a condensation of ammonites, and based on a very complete database). the Thouarsense Chronozone and the beginning of the Thus the observed diversity and palaeobiogeographic Dispansum Chronozone. During the late Dispansum patterns could be only local patterns (i.e. taphonomic Chronozone, species richness slightly decreases and or environmental effects) and not extensible to other remains stable during the late Toarcian. Pseudoradiosa palaeogeographic areas of the southern part of the and Aalensis chronozones share an average level spe- Northwest European Province. However, the observed cies richness compared to Toarcian variations. patterns are coherent with previous observations of The change in specific richness between Margari- other studies (Doyle 1987, 1994; Mariotti et al. 2012), tatus and Serpentinum chronozones is certainly related which indicate slight differences in faunal composition to the Toarcian Oceanic Anoxic Event occurring dur- between the northern parts of the Northwest European ing the Early Toarcian Crisis. The post-crisis recovery Province on the one hand and the southern parts of the of high specific richness in belemnites occurring dur- Northwest European Province plus the Mediterranean ing Bifrons Chronozone is a common pattern already Province on the other. In order to confirm these obser- observed in belemnites (Little & Benton 1995) and vations, it will then be necessary to extend this kind of also in ammonites (Dera et al. 2010). study to further localities in the same palaeogeograph- During the studied period it is also possible to ic region, or others in the Mediterranean Province to observe variations at the generic scale. The most rel- gain a better understanding of the palaeobiogeography evant turnover concerns the Pliensbachian and the of the belemnites. earliest Toarcian with the disappearance of all the Pliensbachian genera (Passaloteuthis, Parapassalo- teuthis, Pseudohastites, and Pleurobelus) and the rise Acknowledgements of an Acrocoelites-dominated population. During the We thank Christian Meister (Museum of Natural His- late Toarcian, the faunal renewal is less sudden with tory, Geneva, Switzerland) for his help and the sharing of a progressive demise of early Toarcian genera (Acro- his knowledge on the biostratigraphy of the studied area. coelites, Cuspiteuthis, and Dactyloteuthis) and the ap- The field work has been possible only by the commitment pearance and domination of new taxa (Rhabdobelus, of Guy and Liette Kronz-Wilmes (Luxembourg), Robert Salpingoteuthis). Haas (Luxembourg) and Alain Faber (National Museum of Natural History, Luxembourg). Domenico Mannetta The faunal composition in Tournadous is relative- (Sapienza University, Rome, Italy) prepared the figured thin ly different from those known from England (Doyle sections. Our acknowledgements go also to Oksana Dzyuba 1990, 1992) and South-west Germany (Riegraf 1980; and Peter Doyle for their advices during the review pro- Riegraf et al.1984). In Tournadous, the genera Mega- cess, which allowed to improve our manuscript. This work teuthis and Brevibelus are absent, whereas they are is a contribution by the INTERVIE (INSU) research pro- Belemnites from the Upper Pliensbachian and the Toarcian (Lower Jurassic) of Tournadous 175 gram and by the BioME team of the Biogéosciences labora- stitutt Skrifter, 189: 1-77. tory (Dijon, France). The work of Jean-Daniel Pinard is Elmi, S., Rulleau, L., Gabilly, J. & Mouterde, R. (1997): supported by the Fonds National de la Recherche (Luxem- Toarcien. – In: Cariou, É. & Hantzpergue, P. (Eds.): bourg) Biostratigraphie du Jurassique ouest-européen et médi- terranéen: zonations parallèles et distribution des in- vertébrés et microfossiles. – Bulletin du Centre de Re- References cherches Elf Exploration et Production, Mémoires, 17: 25-36. Bather, F.A. (1888): Professor Blake and shell-growth in Fischer, J.C. 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