# New sharks and rays from the Cenomanian and Turonian of Charentes, France

Romain Vullo, Henri Cappetta, Didier Neraudeau

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## To cite this version:

Romain Vullo, Henri Cappetta, Didier Nerauvre. New sharks and rays from the Cenomanian and Turonian of Charentes, France. Acta Palaeontologica Polonica, 2007, 52 (1), pp.99-116. hal-00139606

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# New sharks and rays from the Cenomanian and Turonian of Charentes, France

ROMAIN VULLO, HENRI CAPPETTA, and DIDIER NÉRAUDEAU

<figure><img src="image_3.png" /></figure>

Vullo, R., Cappetta, H., and Néraveau, D. 2007. New sharks and rays from the Cenomanian and Turonian of Charentes, France. *Acta Palaeontologica Polonica* 52 (1): 99-116.

New or so far poorly known neoselachians from the Cenomanian and Turonian of SW France are described. The material studied herein comes from nine localities in the Charentes region, comprising palaeoenvironments ranging from coastal to open marine environments, and consists of two orectolobiforms, six lamniforms, and four rajiforms. The new taxa are *Squalicorax coquandi* sp. nov. and *Rouletia bureaui* gen. et sp. nov. within lamniforms, and *Hamrabatis bernardezi* sp. nov., *Archingeayia sistaci* gen. et sp. nov., and *Engolismaia couillardi* gen. et sp. nov. within rajiforms. New specimens of *Odontaspis rochebrunei* Sauvage, 1880 from the type area allow redescription of this taxon, assigned herein to the genus *Cenocarcharias*. Occurrences of *Squalicorax bahariensis*, *S. cf. intermedius*, and *Archaeolamna* sp., previously unrecorded from this region, and *Almascyllium*, a genus generally described from younger strata, are also noted, and improve knowledge of mid-Cretaceous selachian faunas from Western Europe.

**Key words:** Chondrichthyes, Neoselachii, Orectolobiformes, Lamniformes, Rajiformes, Cretaceous, France.

Romain Vullo [romain.vullo@univ-rennes1.fr] and Didier Néraveau [didier.nerauveau@univ-rennes1.fr], Laboratoire de Paléontologie, Géosciences, UMR 6118, Université de Rennes I, 263 avenue du Général Leclerc, 35042 Rennes cedex, France;

Henri Cappetta [cappetta@isem.univ-montp2.fr], Laboratoire de Paléontologie, UMR 5554, Institut des Sciences de l'Evolution, Université de Montpellier II - Sciences et Techniques du Languedoc, Cc 064, Place Eugène Bataillon, 34095 Montpellier cedex 5, France.

## Introduction

The Cenomanian deposits of the Charentes region (administrative departments of Charente and Charente-Maritime), in the northern part of the Aquitaine Basin (SW France), display successions of coastal marine rocks rich in isolated vertebrate remains, especially selachian teeth. As early as the 19th Century, Coquand (1859) and Sauvage (1880) mentioned and described a few specimens, then largely assigned to new species (*Corax elongatus* Coquand, *C. paralelus* Coquand, *C. trapezoidalis* Coquand, *Lamna trigeri* Coquand, *Otodus michoni* Coquand, *Odontaspis rochebrunei* Sauvage, *Galeocerdo tremauxi* Sauvage). The five nominal species created by Coquand, very briefly described, without figures and based on presumed lost material, may constitute nomina dubia. In comparison, the two species established by Sauvage were illustrated: *O. rochebrunei* seems to be a valid taxon, but needing redefinition, whereas *G. tremauxi* might be a junior synonym of *Cretolamna appendiculata* (Agassiz, 1843). Thereafter, in an article enumerating some fish remains from the Mesozoic of Southern France, Priem (1912) recognized with certainty only *C. appendiculata* and *Squalicorax falcatus* (Agassiz, 1843). Since the works of these former authors, very few studies have dealt with the selachians from the Cenomanian of Charentes. Recently, Landemaine (1991), in an article

devoted to the Late Cretaceous elasmobranchs from several localities of SW France, established for the site of “Les Renardières”, near to Tonnay-Charente (Charente-Maritime), a list of 21 taxa of which seven were new species included in four new genera. Recent systematic study of new or previously ignored deposits has yielded abundant material (Vullo et al. 2003, 2005; Vullo 2005). In addition to the re-description and illustration of *O. rochebrunei* Sauvage, 1880, the new forms of selachians are described here.

*Institutional abbreviation*.—UM, Université de Montpellier II (France).

*Other abbreviations*.—AMA, l'Amas; FBN, Font-de-Benon; FVN, Fouras-Vauban; LMS, Le Mas; MTG, Montagan; PDB, Port-des-Barques; PDI, Le Puits des Insurgés; RND, Les Renardières; TLM, Traslemaine.

## Geological settings

Cenomanian deposits occur largely on the northern edge of the Aquitaine Basin, where they outcrop along a NW-SE axis connecting the Armorican Massif with the Central Massif (Fig. 1A). Within the region, the Cenomanian transgression represents the return to marine conditions after the Late Jurassic-Early Cretaceous regressive period. The various
<figure><img src="image_4.png" /><figcaption>Fig. 1. A. Map of the Charentes region showing the Cenomanian and Turonian outcrops and the geographical position of the localities which have yielded the selachian teeth studied in this paper. B. Stratigraphical positions of the localities.</figcaption></figure>
<table><caption>Table 1. List and relative abundance of selachian taxa from each locality (after Landemaine 1991; Vullo et al. 2003, 2005; Vullo 2005). Abbreviations: A, abundant; C, common; R, rare; Tur., Turonian; AMA, l'Amas; FBN, Font-de-Benon; FVN, Fouras-Vauban; LMS, Le Mas; MTG, Montagan; PDB, Port-des-Barques; PDI, Le Puits des Insurgés; RND, Les Renardières; TLM, Traslemaine.</caption><thead><tr><th rowspan="2">Taxa / Localities</th><th colspan="5">Lower Cenomanian</th><th colspan="3">Upper Cenomanian</th><th rowspan="2">Tur.<br>PDB</th></tr><tr><th>FDB</th><th>RND</th><th>FVN</th><th>TLM</th><th>MTG</th><th>PDI</th><th>AMA</th><th>LMS</th></tr></thead><tbody><tr><td>Hybodus sp.</td><td>C</td><td>C</td><td>C</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Polyacrodus maiseyi Landemaine, 1991</td><td>—</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Polyacrodus levis (Woodward, 1887)</td><td>—</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Polyacrodus sp.</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Tribodus morlati (Landemaine, 1991)</td><td>A</td><td>A</td><td>C</td><td>C</td><td>C</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Heterodontus paucicarinatus (Landemaine, 1991)</td><td>—</td><td>R</td><td>—</td><td>C</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Cederstroemia sp.</td><td>—</td><td>R</td><td>—</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Almascyllium sp.</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Cantiosyllium spp.</td><td>A</td><td>R</td><td>C</td><td>C</td><td>C</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Chilosyllium sp.</td><td>C</td><td>—</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>“Carcharias” amonensis (Cappetta and Case, 1975)</td><td>A</td><td>A</td><td>A</td><td>A</td><td>A</td><td>R</td><td>C</td><td>—</td><td>—</td></tr><tr><td>Roulletia bureui gen. et sp. nov.</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Scapanorhynchus minimus Landemaine, 1991</td><td>A</td><td>A</td><td>A</td><td>C</td><td>A</td><td>R</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Squalicorax bahariensis (Stromer, 1927)</td><td>—</td><td>R</td><td>—</td><td>—</td><td>—</td><td>R</td><td>—</td><td>R</td><td>—</td></tr><tr><td>Squalicorax coquandi sp. nov.</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>R</td></tr><tr><td>Squalicorax cf. intermedius Glückman in Glückman and Shvazhaite, 1971</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>R</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Squalicorax sp.</td><td>R</td><td>C</td><td>C</td><td>R</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Cenocarcharias tenuiplicatus (Cappetta and Case, 1975)</td><td>R</td><td>R</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Cenocarcharias rochebrunei (Sauvage, 1880)</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>R</td><td>R</td><td>R</td><td>—</td></tr><tr><td>Cretodus semiplicatus (Münster in Agassiz, 1843)</td><td>R</td><td>C</td><td>C</td><td>R</td><td>R</td><td>—</td><td>R</td><td>R</td><td>—</td></tr><tr><td>Archaeolamna sp.</td><td>—</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Protolamna compressidens (Herman, 1977)</td><td>—</td><td>C</td><td>R</td><td>—</td><td>R</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Protolamna sp.</td><td>—</td><td>R</td><td>R</td><td>R</td><td>R</td><td>—</td><td>R</td><td>R</td><td>—</td></tr><tr><td>Cretolamna appendiculata (Agassiz, 1843)</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>R</td><td>R</td><td>R</td><td>R</td></tr><tr><td>Protoscyliorhinus magnus Landemaine, 1991</td><td>R</td><td>R</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Pseudohypolophus mcnultyi (Thurmond, 1971)</td><td>A</td><td>C</td><td>A</td><td>R</td><td>C</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Turoniabatis cappettai Landemaine, 1991</td><td>C</td><td>R</td><td>C</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Onchopristis dunklei McNulty and Slaughter, 1962</td><td>R</td><td>C</td><td>C</td><td>—</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Ptychotrygonoides pouiti Landemaine, 1991</td><td>R</td><td>C</td><td>C</td><td>C</td><td>C</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Ptychotrygon spp.</td><td>A</td><td>R</td><td>C</td><td>R</td><td>C</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Sclerorhynchid-like indeterminate genus and species</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>Archingeayia sistaci gen. et sp. nov.</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Engolismaia couillardi gen. et sp. nov.</td><td>—</td><td>—</td><td>—</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Hamrabatis bernardezi sp. nov.</td><td>R</td><td>—</td><td>C</td><td>R</td><td>C</td><td>—</td><td>R</td><td>—</td><td>—</td></tr><tr><td>“Rhinobatos” sp.</td><td>—</td><td>—</td><td>R</td><td>R</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td></tr><tr><td>Squatirhina sp.</td><td>—</td><td>—</td><td>—</td><td>R</td><td>—</td><td>—</td><td>—</td><td>—</td><td>—</td></tr></tbody></table>

localities treated here are dated from the basal Cenomanian to the basal Turonian (Fig. 1B). The first facies of the series correspond to very paralic, coastal environments (littoral, lagoons, mangroves, estuaries) (Néraudeau et al. 2002; Vullo et al. 2003) changing to more distal and pelagic environments (outer shelf) at the end of the stage (Moreau 1996). The Cenomanian deposits of the Charentes area are divided into seven lithological units (A to G), themselves subdivided into subunits (e.g., subunits B1, B2, and B3 for unit B), each subunit consisting of one or several lithological levels (e.g., B2gl for the glauconitic upper part of the subunit B2) (Néraudeau et al. 1997).

*Locality 1.*—“Font-de-Benon” quarry at Archingeay – Les Nouillers, Charente-Maritime (Lower Cenomanian; subunit B1, lithological level B1cs): this locality, originally exploited and studied for its levels with insect-bearing amber (Néraudeau et al. 2002), contains in the higher part of the section a condensed sandy layer (60 cm thick), very rich in vertebrate microremains and yielding a diverse invertebrate fauna (orbitolinids, oysters, echinoids) (Vullo et al. 2003). This level, located towards the top of the section, corresponds to the very first clearly marine siliciclastic facies deposited at the beginning of the Cenomanian transgression.
*Locality 2.*—“Les Renardières” quarry at Tonnay-Charente, Charente-Maritime (Lower Cenomanian; subunit B1, lithological level B1s): within a cutting now landscaped, this rich vertebrate microremain deposit corresponds to a 40 to 50 cm thick condensed bed, consisting of phosphatic silty sands. The stratigraphy and the palaeontological characteristics of that locality, sometimes also mentioned in the literature as “Lussant”, have recently been described by Néraveau et al. (2005), after the selachian fauna had been previously studied by Landemaine (1991).

*Locality 3.*—“Vauban” cliff at Fouras, Charente-Maritime (Lower Cenomanian; subunit B2, lithological level B2ms): this locality occurs as a condensed level deposited on a hard ground marking the top of the subunit B1. It comprises 50 cm thick sandy clayey lenses bearing many gravels, encrusting algal concretions, oysters, echinoderms, bone pebbles, and small limonitized wood fragments. The vertebrate remains accompanying the selachian teeth are represented by actinopterygians (pycnodontiforms, *Enchodus* sp.), chelonians, crocodilians, pterosaurs, and dinosaurs (Vullo et al. 2005).

*Locality 4.*—“Traslemaine” at Champniers, Charente (Lower Cenomanian; subunit B2, lithological levels B2ms–B2gl): this site was a temporary section exposed in 2004 by the construction of a new road and is now landscaped. There was here no real distinction between lithological levels B2ms and B2gl. The vertebrate-bearing layer corresponded to an alternation of shelly glauconitic sands and sandstones (50 cm to 3 m thick). The fauna is rather similar to that recovered from Fouras-Vauban.

*Locality 5.*—“Montagan” at Mainxe, Charente (Lower Cenomanian; subunit B2, lithological levels B2ms–B2gl): this site corresponds to a temporary exposure caused in 2005 by the construction of a new road, just south of the city of Jarnac. The glauconitic subunit B2 is lenticular (0 to 4 m thick) and consists of very fine green sands and of an alternation of millimetres-thick clay lenses. Vertebrates and wood microremains are locally highly concentrated. Invertebrate remains are rare and comprise poorly preserved fine shell fragments.

*Locality 6.*—“Le Puits des Insurgés” at Madame Island, Charente-Maritime (Upper Cenomanian; subunits C4–D, lithological levels C4ms–Dm): the cliff of this small island exposes at this place a mound of silty marls (2 m thick), basically indurated (C4ms), and resting in discontinuity on the top of the Middle Cenomanian. In the Angoulême region, this transgressive sequence shows an appreciably different facies, described below. As regards the vertebrate fauna, Madame Island and Angoulême formations have numerous taxa in common but differ from each other in respective proportions of the various groups. The selachian teeth, relatively rare at Madame Island, are associated with common pycnodontiform fish remains as well as vertebrae of a marine aigialosaur varanoid, *Carentonosaurus mineaui* (Rage and Néraveau, 2004).

*Locality 7.*—“L’Amas” quarry at Roullet–Saint-Estèphe, Charente (Upper Cenomanian; subunit D, lithological level Dm):

this layer, originally called “argiles tégulines” by Coquand (1859), is exposed in numerous cement quarries just south of Angoulême. In Angoumois, this unit shows a particular facies characterized by distinctive bluish clay deposits, 5 metres thick, slightly silty-sandy at their base and very rich in oysters (*Cerastostreon flabellatum*). The selachian teeth were collected throughout the whole thickness of the layer.

*Locality 8.*—“Le Mas” quarry at La Couronne, Charente (Upper Cenomanian; subunit D, lithological level Dm): localized close to “l’Amas”, this quarry shows the same levels and facies, and has yielded a similar fauna (see Table 1). Among the vertebrate fauna, pycnodont fish are predominant here.

*Locality 9.*—Port-des-Barques, Charente-Maritime (basal Turonian; subunit T1, lithological level T1m): visible in the northern cliff of the village, the layer that has yielded the unique specimen of *Squalicorax* described below is 1.5 m above the base of a series of beige marls marking the maximum flooding of the Cenomanian–Turonian boundary. The macrofauna is very poor and not very diverse, but contains rare Upper Cenomanian ammonites (*Metoicoceras geslinianum*) in the underlying marls of the subunit G2 (lithological level G2m). Since the publication of the synthetic section by Néraveau et al. (1997), the discovery of a few Lower Turonian ammonites (*Pseudo-tissotia* sp. gr. “*P. inopinata*” Kennedy and Bayliss, 1977) (Philippe Courville personal communication 2004) only one metre above the base of the marls allows us to recognize a slightly lower position of the Cenomanian–Turonian boundary. Consequently, it is here presumed that the tooth has been collected in the earliest Turonian beds.

## Material and methods

Most teeth examined for this study were obtained between 2002 and 2005 using bulk sampling and screen-washing method (sieve sizes used: 0.35–0.71–1 mm), especially for the condensed sandy layers of the Early Cenomanian localities (about 500 kg to 1 t of sediment per locality). The larger teeth, mainly from the Late Cenomanian and Turonian localities, have been surface-collected, mostly since 2000. All numbered and figured specimens are housed in the palaeontological collections of the University of Montpellier II, France. Unnumbered material is housed in the University of Rennes I. The tooth terminology follows that of Cappetta (1987).

## Systematic palaeontology

- Class Chondrichthyes Huxley, 1880

- Subclass Elasmobranchii Bonaparte, 1838

- Cohort Euselachii Hay, 1902

- Subcohort Neoselachii Compagno, 1977

- Order Orectolobiformes Applegate, 1972
<figure><img src="image_5.png" /><figcaption>Fig. 2. Teeth of the hemiscylliid *Almascyllium* Cappetta, 1980a, the ginglymostomatid *Cantioscyllium* Woodward, 1889, the cretoxyrhinid *Archaeolamna* Siverson, 1992 and the anacoracid *Squalicorax* Whitley, 1939 from the Cenomanian and Turonian of Charentes, SW France. **A. Almascyllium** sp., Font-de-Benon, Lower Cenomanian; labial view of UM FBN 1. **B. Cantioscyllium** cf. decipiens Woodward, 1889, l'Amas, Upper Cenomanian; labial (B₁), and occlusal (B₂) views of UM AMA 1. **C. Archaeolamna** sp., l'Amas, Upper Cenomanian; labial (C₁), and lingual (C₂) views of UM AMA 2. **D. Squalicorax baharijensis** Stromer, 1927, Le Mas, Upper Cenomanian; labial (D₁), and lingual (D₂) views of UM LMS 1. **E. Holotype of Squalicorax coquandi** sp. nov., Port-des-Barques, basal Turonian; labial (E₁), distal (E₂), and lingual (E₃) views of UM PDB 1. **F. Squalicorax** cf. *intermedius* Glückman in Glückman and Shvazhaite, 1971, Le Puits des Insurgés, Upper Cenomanian; labial (F₁), mesial (F₂), and lingual (F₃) views of UM PDI 1.</figcaption></figure>
Family Hemiscylliidae Gill, 1862

Genus *Almascyllium* Cappetta, 1980a

*Type species:* *Chiloscyllium cheikheliasi* Signeux, 1949 from the Santonian of Lebanon.

*Almascyllium* sp.

Fig. 2A.

*Material*.—A single isolated tooth crown (UM FBN 1) from Font-de-Benon, Archingeay-Les Nouillers, Lower Cenomanian, base of subunit B1.

*Description*.—The unique specimen corresponds to a lateral tooth devoid of its root. The crown displays a rather slim and pointed cusp, with clearly concave edges in labial view. We observe a pair of relatively rounded lateral cusplets. The distal one is at right angle to the cusp, whereas the smaller mesial one is separated from the main cusp by a slight notch. The basal edge of the apron is damaged. The ornamentation is reduced to five spaced labial folds and not exceeding the height of the cusplets. The three central folds follow the alignment of the cusp whereas each lateral fold is directed towards the respective cusplet.

*Remarks*.—This tooth is morphologically very close to that of *Almascyllium cheikheliasi* (Santonian) illustrated by Cappetta (1980a: 120, fig. 24a–c) in his study on the selachians from the Cretaceous of Lebanon. It differs only from this one by its relatively larger mesial cusplet, as well as by its lower and twice less numerous folds. Bernardez (2002) assigned with doubt a few teeth from the Lower Turonian of the Northern Spain to the genus *Almascyllium*. Those teeth, devoid of lateral cusplets and showing a practically smooth labial face, clearly differ from the tooth here described. Although the specimen UM FBN 1 might be a parasymphyseal or a lateral tooth of *Cantioscyllium* displaying a unusual morphology, like those described by Cappetta and Case (1999: pl. 15: 12–?13) from the Upper Cenomanian of Texas, we assign it to *Almascyllium* sp. because of its rather concave labial face and its salient basal crests parallel to the labial edge of the crown.

Family Ginglymostomatidae Gill, 1862

Genus *Cantioscyllium* Woodward, 1889

*Type species:* *Cantioscyllium decipiens* from the Cenomanian of England.

*Cantioscyllium* cf. *decipiens* Woodward, 1889

Fig. 2B.

*Material*.—A single isolated tooth (UM AMA 1) from l'Amas, Roullet-Saint-Estèphe, Upper Cenomanian D.

*Description*.—This is a complete lateral tooth, broader than high. The triangular main cusp bears on its labial face a few enameloid folds. They are not very strong, and are restricted to the basal half of the crown. The apron is very broad, not prominent, and slightly bifid. There are two pairs of small, blunted, lateral cusplets (damaged on the mesial side). In occlusal view, the medio-lingual protuberance is not very

well developed. The hemiaulacorhize root displays one pair of large margino-lingual foramina.

*Remarks*.—Teeth of *Cantioscyllium* are relatively common in the Lower Cenomanian of Charentes, and even can be very abundant locally, like in the sands of the “Font-de-Benon” quarry at Archingeay-Les Nouillers (Vullo et al. 2003). Despite this, the tooth described herein represents the first occurrence of this genus in the Upper Cenomanian of the Charentes region. Although the labial face of the crown is less densely folded than in *C. decipiens*, this tooth is morphologically close to the specimens figured by Cappetta (1973) and Cappetta and Case (1999), and can be tentatively assigned to the type species. Furthermore, a comparison can also be made with very similar teeth, described by Bernardez (2002) as “*Cantioscyllium* sp. B”, from the Upper Cenomanian of northern Spain. Conversely, the specimen UM AMA 1 differs from the two other Upper Cretaceous species, *C. meyeri* Case and Cappetta, 1997 and *C. nessovi* Ward and Averianov, 1999, which have narrower and more folded lateral teeth. It also differs from the Lower Cretaceous species *C. alhaulfi* Kriwet, 1999, which has teeth with a single pair of cusplets and very few labial folds. Finally, our tooth fits well with the type specimen from the Turonian of England described by Woodward (1889) and illustrated in 1911 (pl. 42: 7–11).

Order Lamniformes Berg, 1958

Family Cretoxyrhinidae Glückman, 1958

Actually, the extent and the content of this family are rather imprecise and subject to discussion. So, a serious revision of this family is needed, but out of the scope of the present article. This problem will be treated by one of us (HC) in a forthcoming study. The assignment of the genus *Archaeolamna* to this family is provisional.

Genus *Archaeolamna* Siverson, 1992

*Type species:* *Odontaspis kopingensis* Davis, 1890 from the Campanian of Sweden.

*Archaeolamna* sp.

Fig. 2C.

*Material*.—One tooth (UM AMA 2) from l'Amas, Roullet-Saint-Estèphe, Upper Cenomanian D; a few unnumbered incomplete teeth from Les Renardières, Tonnay-Charente, Lower Cenomanian B1, and from Vauban cliff, Fouras and Traslemaine, Champniers, Lower Cenomanian B2.

*Description*.—The illustrated lateral tooth (UM AMA 2) shows a relatively narrow cusp curved toward the commissure. The lingual face of the cusp is convex, and the labial face is flat. Both faces are smooth. There is one pair of strong, triangular lateral cusplets. The root is thick and has a well-developed medio-lingual protuberance. The lingual neck is narrow, nearly as much at the centre of the cusp as near the margins. The two lobes are clearly distinct, separated by a rounded basal notch. In the anterior files, the teeth have a more erect cusp and a less laterally expended root.
*Remarks.*—The genus *Archaeolamna* was originally erected by Siverson (1992) for the reassignment of the Davis' species *Odontaspis kopingensis*, from the Campanian of Sweden. This genus occurs commonly from the Albian to the Maastrichtian in numerous regions (Siverson 1992, 1996, 1997; Biddle 1993). A second species, *A. haigi* Siverson, 1996, has been recently described from the Cenomanian of Australia, beside teeth tentatively referred to *A. kopingensis* (Siverson 1996). Conversely to *A. kopingensis*, *A. haigi* have lingually folded teeth. Zhelezko (1990) has described three species and subspecies, *Archaeolamna aduncata*, *A. aduncata suberecta*, and *A. arcuata orica* from the Santonian of western Kazakhstan, under the generic name of *Protolamna* Cappetta, 1980c. Due to the poor quality of the Zhelezko's illustrations, it is difficult to evaluate the validity and mainly the morphological differences between species. According to Mikael Siverson (1996, personal communication 2006), this material entirely corresponds to the type species *A. kopingensis*. The specimen UM AMA 2 is also similar to the lateral teeth of *Dwardius* Siverson, 1999, but it cannot be assigned to this monospecific genus owing to its relatively slenderer cusp and its narrower neck at the centre of the cusp (see Siverson 1999). In *Dwardius woodwardi* (Herman, 1977), the neck is indeed much more developed below the cusp than marginally. Due to the scarcity of the Charentes material, it is here considered preferable to leave the specimens in open nomenclature, as *Archaeolamna* sp.

Family Anacoracidae Casier, 1947

Genus *Squalicorax* Whitley, 1939

*Type species:* *Corax pristodontus* Agassiz, 1843 from the Maastrichtian of the Netherlands.

*Squalicorax baharijensis* (Stromer, 1927)

Fig. 2D.

1927 *Corax baharijensis* Stromer, 1927: 5, pl. 1: 25–27.

1969 *Squalicorax kaupi*; Bilelo 1969: 342, text-fig. 2l-o (?p, ?q).

1974 *Squalicorax baharijensis*; Slaughter and Thurmond 1974: 32, figs. 2b, c.

1974 *Squalicorax curvatus*; Meyer 1974: 298, 300, text-fig. 89.

1989 *Squalicorax baharijensis*; Werner 1989: 35, text-fig. 13, pl. 14: 1–9, pl. 15: 1–3.

1993 *Squalicorax curvatus*; Welton and Farish 1993: 116, figs. 1, 2, and 4 (?fig. 3).

1999 *Squalicorax baharijensis*; Cappetta and Case 1999: 12, pl. 3: 6–9.

2001 *Squalicorax baharijensis*; Case 2001: 106, pl. 2: 3.

2002 *Squalicorax baharijensis*; Bernardez 2002: 279, pls. 42, 43. Unpublished PhD.

*Material.*—Four teeth from Le Puits des Insurgés, Madame Island, l'Amas, Roullet-Saint-Estèphe and Le Mas, La Courronne (UM LMS 1), Upper Cenomanian C4-D; a few teeth from Les Renardières, Tonnay-Charente, Lower Cenomanian B1.

*Description.*—The most characteristic tooth corresponds to an anterior element and is 15.7 mm high (UM LMS 1; Fig. 2D). It is higher than wide, with a crown that is narrow and sharp. The mesial cutting edge shows a regular and distinct convexity,

while the distal one is completely straight. The distal notch and heel are well developed. The cutting edges are finely serrated. The root is rather thick, with the distal lobe more developed than the mesial one and a deep basal notch.

*Remarks.*—This rather large species of *Squalicorax*, restricted to the Cenomanian, displays a broad geographical distribution. It occurs in Egypt (Werner 1989), in North America (Texas, Minnesota) (Cappetta and Case 1999; Case 2001) and in Europe, where it has been recently described from Spain (Bernardez 2002). Its discovery in Charentes is thus not surprising. This species was not recognized by Landemaine (1991) among the *Squalicorax* teeth from the Lower Cenomanian of Les Renardières, all assigned (without illustrations) by this author to *S. falcatus*. Cappetta and Case (1999) and Antunes and Cappetta (2002) noted that the status of the species *S. falcatus* is poorly defined, and Landemaine's assignment thus remains doubtful. However, we collected a few teeth that might correspond to *S. baharijensis*. It is interesting to note that *S. baharijensis* is always accompanied by "*Carcharias* amonensis (Cappetta and Case, 1975), an abundant species occurring in all the Cenomanian localities studied in Charentes (Landemaine 1991; Vullo et al. 2003, 2005; Vullo 2005; Table 1). This association is characteristic of the peri-Tethyan and North American Cenomanian shallow water deposits.

*Squalicorax coquandi* sp. nov.

Fig. 2E.

*Holotype:* Specimen UM PDB 1.

*Derivation of the name:* Species named in memory of Henri Coquand (1811–1881), for his work on the palaeontology of the Cretaceous of Charentes (1859).

*Type locality:* Port-des-Barques, Charente-Maritime.

*Type horizon:* Lowermost Turonian, base of subunit T1, lithological unit T1m.

*Material.*—One complete tooth. Only known from the type locality.

*Diagnosis.*—This large-sized species of *Squalicorax* is easily distinguishable from all other species of the genus by its tooth morphology, characterized by an acute, slim cusp showing subrectilinear edges (according to the single available specimen). The cutting edges are very finely and regularly serrated. There is a clear and rounded distal notch.

*Description.*—The single but perfectly preserved specimen (UM PDB 1; Fig. 2E) is 22 mm high and probably corresponds to an antero-lateral tooth, slightly broader than high. This tooth has a narrow, slim and acute cusp, bent toward the rear. The edges are practically straight: the mesial one is very slightly convex and the distal one very slightly sigmoid. The distal notch is rather open (110°) and is both clear and slightly rounded. The distal heel, well developed, is long, oblique, and almost rectilinear. The cutting edge serrations are very fine and very regular. The basilo-labial bulge of the crown is not very prominent and disappears laterally. The root is rather high and not very thick, with lobes separated by
a broad median concavity. It displays a series of foramina under the labial bulge of the crown.

*Remarks*.—It is sure that the definition of a new species based on a single tooth can be discussed and criticized. However, it is important to note that the species of *Squalicorax* are generally characterized by a weak gradient monognathic and dignathic heterodonty (see Welton and Farish 1993). Thanks to this relative homodonty, one tooth can be enough to display the diagnostic dental features of a species. In this precise case, the holotype is perfectly preserved and clearly different from all other previously described species. A second specimen, unfortunately lost, has also been studied. This large-sized species seems to be very rare. Collecting additional specimens may take a lot of time. So, it seems to us that the description of this new taxon is justified.

By its tooth morphology, *Squalicorax coquandi* sp. nov. differs clearly from the other described species of the genus. However, Antunes and Cappetta (2002: pl. 7: 6–8) have figured, under the name of *Squalicorax* sp., some teeth from the Upper Turonian of Iembe (Angola) showing similarities with *S. coquandi* sp. nov. (narrow and acute cusp, subrectilinear edges). These teeth, however, differ in their smaller size and in having a more acute and angular distal notch. The large size of teeth of *S. pelagicus* sp. nov. is unlike that of other *Squalicorax* of earliest Turonian age. The analogy existing between the teeth of *Squalicorax* and the upper teeth of some large pelagic species of *Carcharhinus* Blainville, 1816 was underlined by Cappetta and Case (1999). If the teeth of *S. falcatus*, with a broader and more robust crown, can be compared for example with those of *C. longimanus* Poey, 1861, *S. coquandi* sp. nov. reminds by its more sharp teeth another oceanic species, *C. falciformis* Müller and Henle, 1841. It also reminds the tooth morphology of the genus *Sphyrna* Rafinesque, 1810.

*Squalicorax cf. intermedius* Glückman in Glückman and Shvazhaite, 1971

Fig. 2F.

*Material*.—A single isolated tooth (UM PDI 1) from Le Puits des Insurgés, Madame Island, Upper Cenomanian, unit D, lithological level Dm.

*Description*.—This tooth is broader than high and rather thick. The cusp is sharp and distally inclined at an angle of about 45°. The mesial cutting edge is relatively straight, with a light concave sinuosity on its lower third. The distal cutting edge is rectilinear and oblique. The distal heel is not very high and slightly convex. The cutting edges are rather strongly but regularly serrated. The distal notch makes an angle of about 90°. In profile view, the tooth is clearly concave labially. The labial face of the crown overhangs the root by a strong and salient bulge cut out by some irregular foraminae that are larger medially. The root is thick and very well separated lingually from the crown; its lingual face shows three irregularly placed foramina.

*Remarks*.—By its size and morphology, our specimen can be related to the *Squalicorax falcatus* "species-group". This "group" probably contains numerous undescribed species

and requires revision, out of the scope of this article. One species has been described from the Upper Turonian of western Kazakhstan, *S. intermedius* Glückman in Glückman and Shvazhaite, 1971. The original illustration is rather poor but a dozen of well preserved specimens from Kazakhstan have been studied by one of us (HC). Our specimen shows a dental morphology that is similar to *S. intermedius*, especially in respect to the basal labial bulge of the crown. In *S. cf. intermedius*, however, the cutting edges of the cusp, and particularly the distal one, are straight, when they are clearly convex in *S. intermedius*.

Family Odontaspididae Müller and Henle, 1839
Genus *Roulletia* nov.

*Type species:* *Roulletia bureui* sp. nov.

*Derivation of the name:* From the name of the type locality.

*Diagnosis and comparisons*.—Small Odontaspididae, characterized by rather bulbous teeth with a cusp showing a strong labial bulge. The lingual and labial faces are completely smooth. There is only one pair of divergent, very small sized, sometimes incipient, lateral cusplets. An additional minute distal cusplet can develop. The mesial edge is slightly concave in antero-lateral teeth. The teeth of this genus have a more bulbous labial face and one pair of proportionally smaller cusplets than in other odontaspidids. They are also distinguished from teeth of “*Carcharias*” *amonensis*, the morphologically closest taxon, by a thicker crown and root (labio-lingually compressed in “*C.*” *amonensis*), and in having fewer and smaller lateral cusplets (generally two pairs in “*C.*” *amonensis*). The teeth of *Roulletia* gen. nov. mainly differ from those of *Cenocarcharias* Cappetta and Case, 1999 by a smooth crown. From the teeth of the cretoxyrhinid? *Dallasiella* Cappetta and Case, 1999 they are distinguished by their weaker lateral cusplets and their clear, well developed nutrient groove on the lingual protuberance of the root.

*Remarks*.—Relatively few odontaspidid genera have been reported in Cretaceous deposits: *Carcharias* Rafinesque, 1810, *Cenocarcharias* Cappetta and Case, 1999, *Hispidaspis* Sokolov, 1978, *Johnlongia* Siverson, 1996, and *Odontaspis* Agassiz, 1838. The genera *Cenocarcharias* and *Hispidaspis* have teeth whose base of the cusp is strongly folded labially, and even lingually for the first one. The teeth of *Roulletia* gen. nov. have a completely smooth enamel and so can be easily separated from *Cenocarcharias* and *Hispidaspis*. *Roulletia* gen. nov. has teeth with cutting edges which are in continuity between the cusp and the cusplets, that separates it from the genus *Odontaspis* in which the cutting edges do not reach the base of the cusp nor of the cusplets, mainly in the anterior files. Moreover, the teeth of *Odontaspis* have sharp and high cusplets, often duplicated, that is not the case of *Roulletia* gen. nov. in which they are simple and low. The teeth of the genus *Johnlongia* have a very particular morphology, mainly in the anterior files whose root shows a very salient lingual protuberance, much more developed than in *Roulletia* gen. nov. In the lateral files, the cusplets are
<figure><img src="image_6.png" /><figcaption>Fig. 3. Teeth of the odontaspidid *Rouletia* gen. nov. from the Cenomanian of Charentes, SW France. *Rouletia bureaui* gen. et sp. nov., l'Amas, Upper Cenomanian. **A.** Specimen UM AMA 3; in labial view. **B.** Specimen UM AMA 4; in labial (B₁), mesial (B₂), and lingual (B₃) views. **C.** Specimen UM AMA 5; in labial (C₁), and lingual (C₂) views. **D.** Holotype, specimen UM AMA 6; in labial (D₁), mesial (D₂), and lingual (D₃) views. **E.** specimen UM AMA 7; in labial (E₁), and lingual (E₂) views. **F.** specimen UM AMA 8; in labial (F₁), mesial (F₂), and lingual (F₃) views. **G.** specimen UM AMA 9; in labial (G₁), and lingual (G₂) views.</figcaption></figure>

sometimes duplicated mesially and the base of the labial face
is often folded. The teeth of *Rouletia* gen. nov. are character-
ized by a not very high cusp compared to the root. In lingual
view, the cusp is indeed practically as high as the root. By
this feature, *Rouletia* gen. nov. is distinguished easily from

*Carcharias* whose teeth, mainly in the anterior files, have a
cusp clearly higher than the root. Moreover, as already noted,
the teeth of *Rouletia* gen. nov. have a completely smooth
enamel, while in *Carcharias* the lingual face of the teeth
bears more or less numerous and marked folds.
Roulletia bureaui gen. et sp. nov.

Fig. 3A–G.

*Derivation of the name:* Species named after Mr. Michel Bureau, the amateur palaeontologist who collected the material.

*Holotype:* Specimen UM AMA 6.

*Paratypes:* Specimens UM AMA 3–5, UM AMA 7–9.

*Type locality:* L'Amas quarry, Roullet-Saint-Estèphe, Charente.

*Type horizon:* Upper Cenomanian, unit D, lithological level Dm.

*Material.*—Seven teeth. Only known from the type locality.

*Diagnosis.*—Same as for genus.

*Description.*—The holotype (UM AMA 6; Fig. 3D) is a lateral tooth, 10 mm high and distally inclined. The smooth crown is relatively narrow, and displays a pair of small cusplets. A second, minute cusplet, can be observed distally. The labial face is slightly swollen. The root is asymmetrical, with a distal lobe stronger than the mesial one. The basilar notch of the root is angular. The lingual protuberance is very developed, and is marked by a deep and relatively broad groove. One of the paratypes (UM AMA 9; Fig. 3G) is an anterior tooth, 9 mm high. It is almost symmetrical, with an erect, sharp cusp, and slightly concave edges. The crown is devoid of ornamentation, and shows convex labial and lingual faces. It bears one pair of very reduced but sharp, acute, marginal cusplets. The root is rather thick, with strong lobes separated by an angular notch. Some specimens, like UM AMA 7 (Fig. 3E), display a rounded root notch.

*Remarks.*—Among odontaspidids, the more reminiscent species is “*Carcharias* *amonensis* (mainly for some teeth of antero-lateral files), whose very distinctive teeth have been found in association with the teeth of *Roulletia bureaui* gen. et sp. nov. in the type locality. So, one could question about the separation of the two taxa at the generic level. Yet, a close study of the material allows us to reject this possibility. In *R. bureaui* gen. et sp. nov., the cutting edge of the crown is in continuity between the cusp and the lateral cusplets, as well in the anterior as in the lateral files. The cusplets are minute and not duplicated, and the root is rather thick with a deep groove. In “*C.*” *amonensis*, in the anterior and antero-lateral files, the teeth lack cutting edges at the base of the cusp and of the cusplets. The teeth of “*C.*” *amonensis* have one to three pairs of lateral cusplets, generally more developed. They are also more labio-lingually compressed, and the crown displays a flatter labial face. The root is more expanded transversally and more flattened labio-lingually (Cappetta and Case 1975; Landemaine 1991). The association of these morphological features allows to separate *R. bureaui* gen. et sp. nov. and “*C.*” *amonensis* at the generic level. It must be also noted that teeth of *R. bureaui* gen. et sp. nov. have never been collected in localities where “*C.*” *amonensis* is abundant, as for instance in the Cenomanian of Texas or in the locality of “Les Renardières”, at Tonnay-Charente. Therefore, *R. bureaui* gen. et sp. nov. cannot be considered as a scarce morphological variation of “*C.*” *amonensis*.

Genus *Cenocarcharias* Cappetta and Case, 1999

*Type species:* *Odontaspis tenuiplicatus* Cappetta and Case, 1975 from the Cenomanian of Texas.

*Cenocarcharias rochebrunei* (Sauvage, 1880)

Fig. 4.

1880 *Odontaspis rochebrunei*, Sauvage 1880: 457, pl. 13: 3.

*Type locality:* L'Amas quarry, Roullet-Saint-Estèphe, Charente.

*Type horizon:* Upper Cenomanian, unit D, lithological level Dm.

*Material.*—Three teeth (UM AMA 10–12) from l'Amas, Roullet-Saint-Estèphe, and one tooth from Le Mas, Upper Cenomanian, unit D.

*Revised diagnosis.*—Odontaspidid with relatively robust, narrow teeth, whose crown displays a strongly folded basal bulge on its labial face, and very reduced ornamentation on lingual face. There is one or two pairs of acutely pointed lateral cusplets, rather linked to the main cusp. The root is rather thick and has a strong lingual protuberance, with a clear groove. This species of *Cenocarcharias* differs from the only other species of the genus, *Cenocarcharias tenuiplicatus* (Cappetta and Case, 1975), by its larger, more robust, and more strongly folded teeth.

*Description.*—An anterior tooth (UM AMA 10; Fig. 4A), 11 mm high, shows a sharp and erect main cusp, relatively narrow and slightly bent distally. The labial face is almost flat whereas the lingual face is convex. The labial face is strongly folded at its base, and the basal ledge is very concave, forming a bulge overhanging the root. The enamel of the lingual face displays a slight granular texture around the root contact. There is one pair of erect marginal cusplets, closely linked to the main cusp. The root shows a strong medio-lingual protuberance bearing a deep groove. A lateral tooth (UM AMA 12; Fig. 4C) has a main cusp very inclined distally, with two pairs of erect lateral cusplets.

*Remarks.*—The original description of this species by Sauvage (1880) was based on only three teeth of the same position (anterior teeth), coming from the “argiles tégulines de la tranchée de Pisani” (locality located on the commune of L'Houmeau-Pontouvre, surroundings of Angoulême). These specimens are currently lost but the discovery of a series of teeth coming from the same area enables us to redefine this taxon. The combination of morphological characters suggests that this species should be referred to the genus *Cenocarcharias* Cappetta and Case, 1999. It is very similar to *Cenocarcharias tenuiplicatus*, a species with slenderer teeth never exceeding 7 or 8 mm (Welton and Farish 1993; Cappetta and Case 1999). However, *Cenocarcharias rochebrunei* may be distinguished from *C. tenuiplicatus* by its larger (up to 11.5 mm), more robust teeth and by its relatively narrower anterior teeth, showing a more pronounced ornamentation on the labial bulge, but a less marked ornamentation on the lingual face of the cusp. It is interesting to note that distinctive teeth of *C. tenuiplicatus* occurs in the Lower Cenomanian of Charentes (Landemaine 1991; Vullo et al. 2005). A tooth from the Cenomanian of Bahariya (Egypt), figured
<figure><img src="image_7.png" /><figcaption>Fig. 4. Teeth of the odontaspidid *Cenocarcharias* Cappetta and Case, 1999 from the Cenomanian of Charentes, SW France. *Cenocarcharias rochebrunei* (Sauvage, 1880), L'Amas, Upper Cenomanian. **A. Neotype**, specimen UM AMA 10; in labial (A₁), mesial (A₂), and lingual (A₃) views. **B. Specimen UM AMA 11; in labial (B₁), distal (B₂), and lingual (B₃) views.** C. Specimen UM AMA 12; in labial (C₁), and lingual (C₂) views.</figcaption></figure>

by Slaughter and Thurmond (1974: fig. 2j) as *Odontaspis* sp. could be referred to *Cenocarcharias* sp.

Superorder Batomorphii Cappetta, 1980b

Order Rajiformes Berg, 1940

Family *incertae sedis*

Genus *Hamrabatis* Cappetta, 1991

*Type species:* *Hamrabatis ornata* Cappetta, 1991 from the Maastrichtian of Egypt.

*Hamrabatis bernardezi* sp. nov.

Fig. 5A–C.

1991 ?*Dasyatis* sp.; Landemaine 1991: 35, pl. 15: 8, 9.

2002 *Hamrabatis sanchezii*; Bernardez 2002: 313, pls. 55, 56, pl. 57: 1. Unpublished Ph.D.

2005 *Hamrabatis* sp.; Vullo et al. 2005: 99, fig. 2.7.

*Derivation of the name:* Species named after Mr. Enrique Bernardez, who originally recognized and described this new species within his unpublished Ph.D. thesis.

*Holotype:* Specimen UM FVN 1.

*Paratypes:* Specimens UM FVN 2, UM MTG 1.

*Type locality:* Vauban cliff, Fouras, Charente-Maritime.

*Type horizon:* Lower Cenomanian, subunit B2, lithological level B2ms.

**Material.**—Two teeth from Font-de-Benon, Archingeay-Les Nouillers, Lower Cenomanian, subunit B1; numerous teeth from Vauban cliff at Fouras, Traslemaine at Champniers and Montagan at Mainxe, Lower Cenomanian, subunit B2; four

teeth from l'Amas, Roullet-Saint-Estèphe, Upper Cenomanian, unit D.

*Diagnosis and comparisons.*—Small species of *Hamrabatis* characterized by its teeth with a rhombic crown in occlusal view, slightly wider than long. Ornamentation consisting of shallow alveoli located at the periphery of the labial face which is rather convex, with a clear but not sharp transverse keel. Lingual face very angular in profile view, with an almost vertical part, below the lingual margin of the labial face. This small-sized species of *Hamrabatis* is easily distinguished from the two other species previously described from Maastrichtian deposits (*Hamrabatis ornata* Cappetta, 1991 from Egypt, and *H. weltoni* Case and Cappetta, 1997 from Texas) by its smaller teeth, displaying a crown less elongated mesio-distally and with a more rhombic outline.

*Description.*—In occlusal view, the holotype (UM FVN 1; Fig. 5A) has a rhombic crown, slightly elongated mesio-distally. A weak transverse keel is developed. The apex of the crown is worn in this specimen. The ornamentation consists of irregular alveoli located on the margins of the occlusal face. The root is nearly half as wide as the crown. In basal view, the root is marked by a deep groove. Some teeth (UM FVN 2 and UM MTG 1; Fig. 5B, C) are narrower in occlusal view, with a crown as wide as long. They are dorso-ventrally less compressed.

*Remarks.*—In his unpublished Ph.D. thesis, Bernardez (2002) recently described a new species of *Hamrabatis* ("*H*.
<figure><img src="image_8.png" /><figcaption>Fig. 5. Teeth of the rajiform *Hamrabatis* Cappetta, 1991 from the Cenomanian of Charentes, SW France. *Hamrabatis bernardezi* sp. nov., Fouras-Vauban (A, B) and Montagan (C), Lower Cenomanian. A. Holotype, specimen UM FVN 1; in occlusal (A₁), basal (A₂), and lingual (A₃) views. B. Specimen UM FVN 2; in occlusal (B₁), lateral (B₂), labial (B₃), and basal (B₄) views. C. Specimen UM MTG 1; in occlusal (C₁), and basal (C₂) views. C₃: enlargement of the root showing the endolithic microborings (ichnotaxon *Abeliella riccioides* Mägdefrau, 1937) near the crown-root junction.</figcaption></figure>

sanchezi”) from abundant material from the Cenomanian of Asturias (northern Spain). The teeth from the Lower Cenomanian of Charentes are similar to the Spanish form although they show a more pronounced ornamentation on both crown faces. Indeed, the Spanish specimens show an enamel very slightly alveolated only at the level of the labial area of the crown. The small differences between these two populations are maybe the result of a preservational bias, and the creation of a further new taxon would not be justified. The original description of this new Cenomanian species by Bernardez (2002) has not been published and “*Hamrabatis sanchezii*” therefore qualifies as a *nomen nudum*, so we herein name this taxon. The genus *Hamrabatis* was originally described from the Maastrichtian of Egypt (Cappetta, 1991), then reported from the contemporary deposits of Texas (Case and Cappetta 1997), Syria (Bardet et al. 2000), and Morocco (Pierre Zennaro personal communication 2003). It also occurs in the

Coniacian of Morocco (HC unpublished data). This Cenomanian species of *Hamrabatis* is distinguished from the Maastrichtian species *Hamrabatis ornata* and *H. weltoni* by its smaller and narrower teeth. This new species was previously assigned to a dasyatid by Landemaine (1991) owing to its tooth morphology, but its histologic tooth type (orthodont) indicates that it corresponds to a rajiform.

## Rajiformes Family *incertae sedis* or Family ?Sclerohynchidae Cappetta, 1974
### Genus *Archingeayia* nov.

*Type species:* *Archingeayia sistaci* sp. nov.

*Derivation of the name:* From the name of the type locality of the type species.

*Diagnosis and comparisons*.—Genus with “*Ptychotrygon*-like” minute teeth with a high, triangular, and clearly cuspi-
date crown in labial (or lingual) view; ornamentation absent or very reduced labially, corresponding generally to a basal marginal ridge; a simple medial vertical labial fold can be developed, bifurcating downward and thus delimiting the apron; one or two pairs of slight vertical lingual folds; relatively strong labio-lingual compression. The root is relatively low, half to third as high as the crown.

Among oral teeth of Sclerorhynchidae, *Ptychotrygon* sensu scicto and closely related forms, *Archingeayia* gen. nov. is firstly well differentiated from the “cross-like crown” group bearing a well developed apron (*Dalpiazia* Chechia-Rispoli, 1933, *Ischyrhiza* Leidy, 1856, *Kiestus* Cappetta and Case, 1999, *Onchopristis* Stromer, 1917, *Pucapristis* Schaeffer, 1963, *Plicatopristis* Cappetta, 1991, *Renpetia* Werner, 1989). It is closer to *Baharipristis* Werner, 1989, *Biropristis* Suarez and Cappetta, 2004, *Borodinopristis* Case, 1987, *Celtipristis* Kriwet, 1999, *Ctenopristis* Arambourg, 1940, *Ganopristis* Arambourg, 1935, *Libanopristis* Cappetta, 1980b, *Micropristis* Cappetta, 1980b, *Ptychotrygon* Jaekel, 1894, *Ptychotrygonoides* Landemaine, 1991, *Sclerorhynchus* Woodward, 1889, and *Texatrygon* Cappetta and Case, 1999. However, teeth of the species of *Ptychotrygon* are generally low cusped, well ornamented and not strongly labio-lingually compressed. *Ptychotrygonoides*, *Sclerorhynchus*, *Ganopristis*, *Biropristis*, and *Borodinopristis* have a radial labial ornamentation. *Celtipristis* shows a strong constriction between the crown and the root. Teeth of *Libanopristis* and *Micropristis* have a rather similar reduced ornamentation but are both relatively low cusped. Teeth of *Texatrygon* have either a completely smooth crown or numerous short vertical labial folds. Anterior teeth of *Ctenopristis* show a more distinct and high cusp. Anterior teeth of *Baharipristis* have a hooked crown strongly bent labio-lingually.

*Archingeayia sistaci* gen. et sp. nov.

Fig. 6A–C.

*Derivation of the name:* Species named after Mr. Paul Sistac, geologist, for his valuable assistance in the field.

*Holotype:* Specimen UM FBN 4.

*Paratypes:* Specimens UM FBN 2–3.

*Type locality:* Font-de-Benon quarry, Archingeay-Les Nouillers, Charente-Maritime, western France.

*Type horizon:* Lower Cenomanian, subunit B1, lithological level B1cs.

*Material*.—Ten teeth. Only known from the type locality.

*Diagnosis*.—Same as for genus.

*Description*.—The holotype (UM FBN 4; Fig. 6C) has a rather high, triangular cusp, slightly worn at its apex. The labial face is smooth, except the presence of a basal horizontal ridge delimiting a bulge. The medio-labial protuberance is not well developed. The lingual face is more steep in profile view and shows three oblique folds not reaching the apex. There is a distinct basal lingual bulge. The uvula above the root notch is rather well differentiated and bears a weak interlocking hollow. The root is relatively low (about one quarter

of the total tooth height). It is slightly narrower than the crown and shows typical triangular lobes in basal view.

*Remarks*.—*Archingeayia* gen. nov. is based on isolated oral teeth. The systematic affinities of this new genus thus remain uncertain because of its unique tooth morphology. In the locality of Archingeay-Les Nouillers, *Archingeayia* gen. nov. occurs in association with *Ptychotrygonoides* and small teeth of *Ptychotrygon* spp. However, the presence of a true Sclerorhynchidae is attested by the discovery of numerous small rostral teeth that seem to belong to a single taxon. For the moment, it is reasonable not to attempt to link these rostral teeth to any oral ones.

Teeth of *Archingeayia* gen. nov. display a particular morphology, with a straight and well cusped triangular crown. However, they are close to some other Cenomanian forms, especially from the Bahariya Formation, Egypt. Werner (1989: pl. 22: 1, 2) incorrectly linked to the rostral teeth of *Onchopristis numidus* some oral teeth showing clear resemblances with teeth of *Archingeayia* gen. nov. (well developed cusp; ornamentation pattern). However, these teeth are characterized by their larger size, their root wider than the crown, and by their cusp bent labio-lingually. The set of teeth described by Werner should be assigned to the genus *Kiestus*, recently defined from the Turonian–Coniacian boundary of Texas (Cappetta and Case 1999). The teeth of *Archingeayia* gen. nov. also show affinities with a few teeth (lateral ones) of *Baharipristis* (Werner 1989: pl. 29: 2). But, as mentioned above in the “Diagnosis and comparisons”, *Baharipristis* shows a strong monognathic heterodonty and remains clearly different from *Archingeayia* gen. nov.

Family, genus, and species indet.

Fig. 7E.

*Material*.—One single tooth (UM AMA 13) from l’Amas, Roullet–Saint-Estèphe, Upper Cenomanian D.

*Description*.—This unique specimen corresponds to a small “*Ptychotrygon*-like” tooth, with a cusped crown. In occlusal view, the medio-labial protuberance is broad and not detached from the general outline. The labial face displays a very reduced ornamentation, with a single vertical ridge linking the apex to the apron. A weak fold is also present along one of the two of the basilo-labial margins. The lingual face is smooth, with a shallow central depression not well delimited. The root is slightly broader than the crown, with a very flat basal face.

*Remarks*.—This tooth represents the only occurrence of a sclerorhynchid-like rajiform in the Upper Cenomanian of Charentes. This contrast with the abundance of teeth of this group in the lower part of the stage (subunits B1 and B2) (Landemaine 1991; Vullo et al. 2003, 2005). The lack of a transverse ridge along the middle of the lingual face separates this specimen from the genus *Ptychotrygon*. It also differs from the genus *Ptychotrygonoides* in lacking radiating labial ridges. A very similar form (small size, cusped crown, same reduced ornamentation) has been described by Ber-
<figure><img src="image_9.png" /><figcaption>Fig. 6. Teeth of the rajiform *Archingeayia* gen. nov. from the Cenomanian of Charentes, SW France. *Archingeayia sistaci* gen. et sp. nov., Font-de-Benon, Lower Cenomanian. **A.** Specimen UM FBN 2; in lingual (A₁), basal (A₂), and lateral (A₃) views. **B.** Specimen UM FBN 3; in lingual (B₁), basal (B₂), and occlusal (B₃) views. **C.** Holotype, specimen UM FBN 4; in lingual (C₁), labial (C₂), occlusal (C₃), and basal (C₄) views.</figcaption></figure>

nardez (2002) from the Middle–Late Cenomanian of north-
ern Spain, for which he proposed a new species, “*Ptycho-
trygonoides lamoldai*” (unpublished, *nomen nudum*). The as-
signment of the Spanish species to this genus, based mainly
on the cuspidate crown, remains doubtful as these teeth are
clearly different from those of the type species, *P. pouiti*
Landemaine, 1991.

Genus *Engolismaia* nov.

Type species: *Engolismaia couillardi* sp. nov.

*Derivation of the name:* Derived from *Engolisma*, the Latin name of the town of Angoulême.

*Diagnosis and comparisons.*—Small rajiform genus, charac-
terized by minute (up to 1 mm wide), cuspidate teeth, of

which the transverse keel develops one to two distinct and
blunt pairs of cusplets, slightly lower than the main cusp. The
rather prominent medio-labial protuberance bears a single
pustule developed from a vertical medial ridge. The root is
relatively high.

*Engolismaia* gen. nov. differs from all other Rajiformes
by its well-cusped crown displaying one or two pairs of
well-developed rounded lateral cusplets, and by the orna-
mentation of the apron consisting of one single pustule. An-
other monospecific sclerohynchid genus, *Baharipristis*, can
also display a pair of lateral cusplets (Werner 1989: pl. 26: 4),
but these cusplets remain very reduced compared to the main
cusp. Moreover, the main cusp of *Baharipristis bastetiae* is
not erect, but clearly inclined labio-lingually.
<figure><img src="image_10.png" /><figcaption>Fig. 7. Teeth of the rajiform *Engolismaia* gen. nov. from the Cenomanian of Charentes, SW France. *Engolismaia couillardi* gen. et sp. nov., Traslemaine, Early Cenomanian. **A.** Holotype, specimen UM TLM 1; in occlusal (A₁), lingual (A₂), basal (A₃), labial (A₄), and lateral (A₅) views. **B.** Specimen UM TLM 2; in occlusal (B₁), lingual (B₂), and lateral (B₃) views. **C.** Specimen UM TLM 3; in occlusal view. **D.** Specimen UM TLM 4; in lingual (D₁), lateral (D₂), and occlusal (D₃) views. **E.** Rajiformes indet., l'Amas, Late Cenomanian; occlusal (E₁), basal (D₂), and lingual (D₃) views of UM AMA 13.</figcaption></figure>
*Engolismaia couillardi* gen. et sp. nov.

Fig. 7A-D.

*Derivation of the name:* Species named after Mr. Alain Couillard, for his valuable assistance in the field.

*Holotype:* Specimen UM TLM 1.

*Paratypes:* Specimen UM TLM 2-4.

*Type locality:* Traslemaine, Champniers near Angoulême, Charente.

*Type horizon:* Lower Cenomanian, subunit B2, lithological level B2gl.

*Material.*—Fifteen teeth from Traslemaine, Champniers, and three teeth from Montagan, Mainxe, Lower Cenomanian, subunit B2.

*Diagnosis.*—Same as for genus.

*Description.*—The holotype (UM TLM 1; Fig. 7A) is a presumed antero-lateral tooth, 0.9 mm wide. In occlusal view, the crown shows a broad, rounded, well-defined apron. The two pairs of cusplets are symmetrically developed on each side of the main cusp. All the cusps have a rounded apex. The basilo-labial ridge is well distinct. The main cusp bears two subvertical folds on its lingual face. The lingual central depression is rather deep. The root, slightly narrower than the crown, displays a flattened basal face, and bears two well-marked foramina on its lingual face. An anterior (or posterior?) tooth (UM TLM 4; Fig. 7D) is narrow, with only one poorly defined lateral cusplet widely united to the base of the main cusp. The other side corresponds to a simple heel. The apron bears a single, oval pustule. The root is relatively high (about one third of the total tooth height).

*Remarks.*—*Engolismaia* gen. nov. is a very characteristic element of the Charentes selachian fauna because of its unusual tooth morphology. It is currently known only to occur in the eastern part of the Charentes region (Montagan and Traslemaine localities) and is restricted to the middle part of the Lower Cenomanian.

Comments on palaeoecology
and palaeobiogeography

Adding to the systematic results previously supplied by Landemaine (1991), the present study increases our knowledge of the selachian faunas from shallow detrital and terrigenous Cenomanian facies of SW France. In addition, a new species of *Squalicorax* is also described from the deeper basal Turonian chalky marls. The selachian species lists are given for the different localities in the Table 1.

Combined with previous works (Landemaine 1991; Vullo
et al. 2003, 2005; Néraudeau et al. 2005), these new data al-
low to refine the palaeoecological and palaeobiogeographi-
cal interpretations for the selachian faunas of this area. The
shallow sandy deposits of the Lower Cenomanian indicate
coastal paralic habitats that seem to have been favourable to
the development of small orectolobiforms and rajiforms. Nu-
merous extant species (e.g., bamboo and nurse sharks) are
known to grow, mate, and live in such environments, like

mangroves (Compagno 2001). By analogy, the existence of
nursery grounds has been suggested by Vullo et al. (2003) for
the earliest Cenomanian of Charentes. Small sclerorhynchid
sawfishes (or closely related forms) are abundant in these
detrital and terrigenous sediments, as it can be also observed
in numerous localities from the Cenomanian of Texas (Wel-
ton and Farish 1993; Cappetta and Case 1999), Spain (Ber-
nardez 2002), and Egypt (Werner 1989). In the Upper Ceno-
manian Bahariya Formation, Egypt, where the selachian
fauna and other vertebrates is very similar to the Charentes
fauna, the palaeobotanical content and the depositional envi-
ronment indicate a *Weichselia*-dominated mangrove (Laco-
vara et al. 2001). Interestingly, this mangrove tree fern oc-
curs in the Early Cenomanian deposits of Charentes (Bernard
Gomez, personal communication 2005), as some gymno-
sperms (*Frenelopsis*, *Glenrosa*) and angiosperms (*Pseudo-
asterophyllites*) characterizing salt marshes (Gomez et al.
2004; Fejfar et al. 2005). All these data show that palaeoeco-
logical conditions existing in Southern Tethys may have ex-
tended northward up to the Aquitaine Basin. The Charentes
selachian fauna differs mainly from Northern European
(England, Belgium, Germany) deeper water assemblages by
the lack of squaloid, palaeospinacid, and sclylorhinid sharks,
common in the Boreal realm (Herman 1977; Müller and
Diedrich 1991; Underwood and Mitchell 1999).

During the Late Cenomanian, deeper and more open ma-
rine environments (“argiles tégulines” and chalky marl fa-
cies) tend to develop in Charentes, with the proliferation of
large lamniforms (e.g., *Squalicorax*) and the local appear-
ance of some cosmopolitan taxa (e.g., *Cretolamna*). This
 contrasts with the strong decrease of the abundance and the
diversity of small benthic forms, like orectolobiforms.

In conclusion, these new data confirm that the selachian
fauna from the Cenomanian of Charentes shares numerous
affinities with those recovered from North America (Texas,
Minnesota) (Cappetta and Case 1999; Case 2001), Southern
Europe (Spain) (Bernardez 2002), and Northern Africa
(Egypt) (Werner 1989), as previously noted by Landemaine
(1991), Cappetta and Case (1999) and Vullo et al. (2005). This
suggests, in combination with sedimentological and syneco-
logical data, similar climatic conditions and depositional envi-
ronments, like warm tropical shallow coastal waters.

Acknowledgements

The authors would like to thank the amateur palaeontologists who ac-
tively participated in the collection of the material or who provided
some specimens: Michel Bureau, Alain Couillard, Jean-Pierre Texier
(Angoulême, France), and Paul Sistac (Toulouse, France). Enrique
Bernardez (Oviedo, Spain) kindly provided Spanish material for com-
parison. Philippe Courville (Géosciences, University of Rennes 1,
Rennes, France) is also thanked for identifying the ammonite from
Port-des-Barques. We are grateful to Vincent Perrichot and Blaise
Videt (both Géosciences, University of Rennes 1, Rennes, France) for
their help concerning the illustrations. The authors wish to thank
Mikael Siverson (Mosman Park, Australia), Charlie J. Underwood
(School of Earth Sciences, Birkbeck College, London, Great Britain), and David J. Ward (School of Earth Sciences, University of Greenwich, Great Britain) for their incisive comments on the manuscript. Contribution ISEM n° 2005-097.

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