Selachians from the type Campanian area (Late Cretaceous), Charentes, western France

Cretaceous Research 26 (2005) 609e632 www.elsevier.com/locate/CretRes

Selachians from the type Campanian area (Late Cretaceous), Charentes, western France Romain Vullo Laboratoire de Pale´ontologie, Ge´osciences, UMR 6118, Universite´ de Rennes 1, 263 avenue du Ge´ne´ral Leclerc, 35042 Rennes cedex, France Received 10 August 2004; accepted in revised form 1 March 2005 Available online 6 September 2005

Abstract The stratotypical Campanian of the northern Aquitaine Basin has yielded a relatively rich and diverse selachian fauna. This fauna consists of 20 taxa, two of which represent new species: Carcharias adneti sp. nov. and Rhombodus carentonensis sp. nov. Additionally, one new combination is proposed: Serratolamna khderii (Zalmout and Mustafa, 2001) comb. nov. The discovery of Rhombodus in the Campanian constitutes the earliest occurrence of the genus and may have widespread biostratigraphical implications. The fauna corresponds to a mixture of Tethyan and Boreal species resulting from the particular palaeogeographical situation of Charentes during the Late Cretaceous. The succession of the selachian faunas from the Middle to Late Campanian shows an association mainly dominated by Boreal species (e.g., Carcharias latus) that becomes progressively influenced by immigrations of typical Tethyan components (e.g., Serratolamna khderii). It provides evidence of palaeogeographical, palaeoenvironmental and palaeoclimatical changes that occurred in the region at the end of the Campanian. Ó 2005 Elsevier Ltd. All rights reserved. Keywords: Neoselachians; Type Campanian; Charente-Maritime; France; New taxa; Palaeobiogeography

1. Introduction The selachians from the Late Cretaceous of the northern Aquitaine Basin have been the object, until now, of very few studies and have thus remained very poorly known. Recently, Landemaine (1991), then Vullo et al. (2003), reported Cenomanian faunas from the surroundings of Rochefort (Charente-Maritime). The former author also announced the presence of seven taxa in the Upper Santonian of Sarlat (Dordogne). Regarding the Campanian, only three authors (Coquand, 1859; Sauvage, 1880; Priem, 1912) have been interested in the remains of selachians from this region. Coquand (1859), after having established the type Campanian within the ‘‘Champagne charentaise’’, briefly described three species from Montignac (Dordogne), ‘‘Otodus marroti’’, ‘‘Lamna petrocoriensis’’ E-mail address: [email protected] 0195-6671/$ - see front matter Ó 2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2005.03.006

and ‘‘Ptychodus pauli’’. These taxa were erected in the absence of any illustrations and could be considered respectively as junior synonyms of Cretolamna appendiculata Agassiz, Cretoxyrhina mantelli (Agassiz) and Ptychodus rugosus Dixon, according to their descriptions. Moreover, the age of this locality, thought by Coquand to be Campanian, is in fact Coniacian (Arnaud, 1887). Priem (1912) recorded two species, ?Scapanorhynchus sp. and Squalicorax pristodontus (Agassiz), from the ‘‘Dordonian’’ (ZUpper Campanian) of Talmont and Meschers, near Royan. Since then, the Campanian selachians of the Charentes region have been essentially ignored for almost a century. Moreover, the previous studies suffer from their age and hence their low objectivity. A few recent studies deal with selachians from the Campanian and Maastrichtian of the southern Aquitaine Basin (Gheerbrant et al., 1997; Cappetta and Odin, 2001) and from the Maastrichtian of the Basque-Cantabrian Basin (Cappetta and

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La Rochelle

FRANCE

A

Rochefort

Saintes Cognac

Royan MESCHERS TALMONT BARZAN

Angoulême Barbezieux

ATL AN TIC

"Champagne charentaise"

OCEAN

Aubeterre

N Locality 0

50 km

Bordeaux Suzac Point

B MESCHERS Arces

Meschers cliffs

G E

0

S

T

I

TALMONT R U

O A

N R

D

E

Y

1 km

Le Caillaud cliff

BARZAN Le Pilou cliff

Fig. 1. Maps showing the localities studied. A, location of the study area within the western part of the Charentes region (west-central France). B, detailed map showing the location of the cliffs examined.

Corral, 1999). These faunas are geographically the nearest to that studied here. The discovery and the description of new material collected from within the type Campanian succession will therefore, allow a better understanding of the faunal composition of, and palaeobiogeographical influences on, this reference area located half-way between the Boreal and the Tethyan realms.

2. Location and geological setting The material discussed herein comes from the cliffs bordering the right bank of the Gironde Estuary, a few kilometres to the south-east of Royan (administrative

department of Charente-Maritime), between Barzan upstream and Meschers downstream (Fig. 1A, B). In this area, estuarine erosion constantly cuts into Campanian deposits, leading to periodical renewal of the sections. This region corresponds to the western extension of the historical Campanian stratotype, defined by Coquand (1857) as a group of beds occurring in the ‘‘Champagne charentaise’’ (southern part of the administrative department of Charente), including the historical section of Aubeterre (van Hinte, 1979; Neumann and Odin, 2001) (Fig. 1A). The type Campanian is generally divided, on the basis of micropalaeontological data (Andreieff and Marionnaud, 1973; Platel, 1977; Neumann et al., 1983), into eight biozones (CIeCVIII) based on associations of

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Localities

First occurrences of index benthic foraminifera

Stratigraphic ranges of ammonites

Depositional environments outer neritic

inner neritic

Meschers

CVII

Synthetic lithostratigraphic section

yellowish chalky limestones with glauconite and oyster lumachelles (P. vesicularis)

Le Caillaud

CIVb

grey marly chalk with silicifications, glauconite, green burrows, silicified beds and oyster lumachelles

Le Pilou

CV

white chalky marly limestones

CIVa

Middle

Campanian

CVI

Upper

CVIII

Stage

Biozonation

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grey chalky limestones with silicified beds yellowish-white marls

CIII

white chalky limestones with glauconite

grey chalky limestones with grey flints bed with grey silicifications

CII CI

Lower

50 m

white chalky limestone

white argillaceous chalky limestone

0

Fig. 2. Biozonation, lithology, index fossils and environments of the type Campanian: data compiled from Neumann et al. (1983), Villier et al. (1997) and Neumann and Odin (2001).

benthic foraminifera (Fig. 2). Thus, the Upper Campanian (CVIeCVIII) is, for example, well characterized in Charentes by the occurrence of Orbitoides media (d’Archiac). Ammonites and belemnites are very rare and generally poorly preserved, not readily allowing any usable zonation based on cephalopods. However, such a biostratigraphy was attempted by Kennedy (1986) and the ranges of the main ammonite species are given in Fig. 2.

Three localities, stratigraphically distributed between the biozones CIV and CVIII, have yielded the material studied herein: (1) Le Pilou cliff, Barzan, Middle Campanian (biozone CIV and lower part of biozone CV); (2) Le Caillaud cliff, Talmont, Middle Campanian (upper part of biozone CIV and biozone CV); (3) Meschers cliffs, Upper Campanian (biozone CVIII). These cliffs are composed of marly limestones, sometimes chalky, with siliceous or glauconitic layers, and are

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often very fossiliferous. Lumachelles of pycnodont oysters [Pycnodonte vesicularis (Lamarck)] occur mainly in the upper part of the series, where they become more and more frequent, notably in biozone CVIII. Depositional environments are within the neritic zone. However, there is a deepening between CI and CV, then a global tendency to shallowing between CV and CVIII, that leads to definitive emergence of the region during the latest Cretaceous (Ne´raudeau and Villier, 1997) (Fig. 2). The material discussed herein was obtained by surface collecting during the past ten years. The technique of washing and sieving was attempted but did not yield selachian material, because of the wide dispersion of the teeth in the sediment (no concentration horizon, high rate of sedimentation). The state of preservation of the teeth is generally good to excellent, although some are devoid of their root. Associated with the teeth, common remains of other vertebrates were also discovered. They consist of an indeterminate chimaeroid, actinopterygians (Anomoeodus, Cimolichthys, Cylindracanthus, Enchodus, Stephanodus), mosasaurids (Plioplatecarpus, Prognathodon) and an indeterminate chelonioid marine turtle. It is interesting to note that this turtle is represented by a fragmentary plastron that shows shark bites like those recently described from the Upper Cretaceous of North America (Schwimmer et al., 1997; Shimada and Hooks, 2004). All numbered selachian specimens studied are housed in the palaeontological collections of the Museum of Natural History of La Rochelle (MHNLR) (CharenteMaritime, France).

3. Systematic palaeontology

Material. Four teeth: PIL 66, 67, 103, CLD 26. Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV and CV). Description. The best preserved tooth (PIL 103; Fig. 3A, B) corresponds to a small anterior tooth, that is broader than it is high and has a labio-lingually compressed sharp cusp. In lateral view, the labial face is concave whereas the lingual face is convex. The cusp is slightly tilted labio-lingually and mesio-distally. In labial view, heels are relatively low and are not clearly separate from the cusp. The apron is narrow and rather pointed. The hemiaulacorhize root is flattened but has a strong medio-lingual protuberance on its upper face. There is a line of marginal foramina on both sides of this protuberance. The basal face of the root is planar and has a sub-rhombic profile. Remarks. These teeth are very close to Squatina hassei, well known from the Campanian and Maastrichtian of Belgium and the Netherlands. However, the identification at the specific level of the teeth of Squatina is difficult (Cappetta, 1987b). Only the discovery of more teeth will allow confirmation of the attribution of the Royannais form to S. hassei. Teeth of Squatina are distinguished from the squatinid-like teeth of the genera Cretorectolobus Case and Cederstroemia Siverson by their less bulky aspect, more distinct cusp, lower heels and the constant hemiaulacorhize stage of the root through time. Superorder: Galeomorphii Compagno, 1973 Order: Heterodontiformes Berg, 1937 Family: Heterodontidae Gray, 1851 Genus Heterodontus Blainville, 1816

The systematics and terminology follow Cappetta (1987b).

Type species. Squalus philippi Schneider, 1801.

Class: Chondrichthyes Huxley, 1850 Subclass: Elasmobranchii Bonaparte, 1838 Subcohorte: Neoselachii Compagno, 1977 Superorder: Squatinomorphii Compagno, 1973 Order: Squatiniformes Buen, 1926 Family: Squatinidae Bonaparte, 1858 Genus Squatina Dume´ril, 1906

Heterodontus cf. rugosus (Agassiz, 1839) Fig. 3C

Type species. Squalus squatina Linnaeus, 1758. Squatina cf. hassei Leriche, 1929 Fig. 3A, B 1929 Squatina hassei Leriche, pp. 206e208, p. 206, textfigs. 1e3. 1964 Squatina hassei Leriche; Albers and Weiler, p. 14, figs. 16e18. 1977 Squatina hassei Leriche; Herman, p. 124, pl. 5, fig. 3.

1839 Acrodus rugosus Agassiz, p. 148, pl. 22, figs. 28, 29. 1964 Heterodontus rugosus (Agassiz); Albers and Weiler, p. 6, figs. 6, 7. 1977 Heterodontus rugosus (Agassiz); Herman, p. 90, pl. 3, fig. 5. Material. 18 teeth: PIL 16e18, 75, 108, CLD 5e9, 37, 38, 45, 54e56, MSH 14, 15. Localities and age. Le Pilou, Le Caillaud and Meschers; Middle Campanian (CIV and CV), Late Campanian (CVIII). Description. An anterior tooth (CLD 8) is rather badly preserved and comprises a massive cuspidate

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crown. There is a pair of very reduced rounded lateral cusplets. The labial face is mesio-distally strongly convex and concave in lateral view. The apron, although damaged, shows an alveoli-ornamented enameloid. This ornamentation is transformed into fine and irregular wrinkles in the first third of the cusp. The remainder of the crown, up to the apex, is completely smooth. The smooth lingual face and the root are very damaged. An antero-lateral tooth (CLD 9) presents a very prominent conical cusp without lateral cusplets. On both sides of this cusp, sub-horizontal lateral expansions are developed, on which the edges are expanded (with only one of the two expansions being preserved). This corresponds to the formation of the transverse crest present on lateral teeth. The apron is reduced. The labial face shows the same ornamentation as on the anterior tooth. The lingual face is completely smooth and has a strong medio-lingual protuberance. On the lateral expansions, the alveoli ornamentation tends to extend lingually beyond the crest. The root is damaged. The lateral teeth, like CLD 7 (Fig. 3C), can reach a rather large size (up to 11 mm in mesio-distal width). The transverse crest is quite marked with a bulge corresponding to a small tubercle, generally worn down by the durophagous activity of these sharks, in its median part. In occlusal view, this crest is sigmoidal. The labial part of the crown always has an alveoliornamented surface. The ornamentation on the lingual part consists of ridges perpendicular to the transverse crest. These folds generally bificate, and can, on certain teeth, be replaced in the basal part of the crown by alveoli. The anaulacorhize roots are generally badly preserved. Remarks. The Royannais teeth are practically indistinguishable from teeth of Heterodontus rugosus from the type Maastrichtian area. H. rugosus is also present in the Campanian and Maastrichtian of Belgium (Herman, 1977) and in the Campanian of Germany (Thies and Mu¨ller, 1993). However, the specific assignment of Heterodontus teeth remains difficult, and this species is also very close to H. lonzeensis Herman (Santonian and Campanian of Belgium, Campanian of Germany) by its ornamentation of the anterior teeth, and to H. havreensis Herman (Campanian of Belgium and Germany) by its lateral teeth. In the Late Senonian, the abundance of Heterodontus in the Boreal region (Anglo-Franco-Belgian Basin, Germany, Sweden) contrasts with its absence from the occidental Tethyan region (Spain, Morocco). Its discovery in the Royannais, which is the southernmost locality in Europe to date, extends the limits of its distribution. Heterodontus also occurs in the south-eastern Tethyan province (Early Maastrichtian of Israel) with presumed deep water taxa (Squaliformes) and other Boreal genera (Paraorthacodus, Pseudocorax) (Lewy and Cappetta, 1989).

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Order: Orectolobiformes Applegate, 1972 Family: Ginglymostomatidae Gill, 1862 Genus Plicatoscyllium Case and Cappetta, 1997 Type species. Plicatoscyllium deramei Case and Cappetta, 1997. Plicatoscyllium cf. globidens (Cappetta and Case, 1975) Fig. 3D, E

1975 Ginglymostoma globidens Cappetta and Case, p. 12, pl. 9, figs. 24, 25, text-fig. 6. 1984 Ginglymostoma globidens Cappetta and Case; Lauginiger, p. 142, pl. 2, figs. 16, 17. 1987 Ginglymostoma globidens Cappetta and Case; Case, p. 17, pl. 5, figs. 6e8. 1988 Ginglymostoma globidens Cappetta and Case; Case and Schwimmer, p. 294, fig. 4.13e16. ?1991 Ginglymostoma globidens Cappetta and Case; Case, p. 2, pl. 1, fig. 1. 2004 Plicatoscyllium globidens (Cappetta and Case); Case and Cappetta. Material. Two teeth: PIL 116, CLD 64. Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV). Description. The only complete tooth (PIL 116; Fig. 3D, E) is rather badly preserved and corresponds to a lateral tooth, which is symmetrical and broader than high. The main cusp, whose apex is strongly worn, is flanked by a pair of relatively well detached cusplets, at the base of which is a second, incipient, pair. The labial face has a broad apron, which is not very prominent. The lower part of this face is spread out labially, which forms in lateral view a strong concavity. The basilo-labial bulge shows a distinct thickening at the apron level. The ornamentation of the crown is very reduced and is limited to two or three folds on the upper part of the labial face. The wear of the main cusp does not allow a more precise description of the ornamentation. The medio-lingual protuberance of the crown and the root are poorly preserved. Remarks. Initially assigned to the genus Ginglymostoma, this species was placed in Plicatoscyllium Case and Cappetta, 1997 by Case and Cappetta (2004), as previously suggested by Noubhani and Cappetta (1997). Indeed, this genus is characterized by its small teeth, an ornamentation that is limited to some more or less strong folds between the median and apical parts of the labial face, and by the restricted number of pairs of cusplets (1e3 pairs). P. globidens, which was previously known only from the Upper Campanian and Lower

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Fig. 3. A, B, Squatina cf. hassei Leriche, lateral tooth in labial (A) and lateral (B) views, PIL 103. C, Heterodontus cf. rugosus (Agassiz), lateral tooth in occlusal view, CLD 7. D, E, Plicatoscyllium cf. globidens (Cappetta and Case), lateral tooth in labial (D) and occlusal (E) views, PIL 116. F, Cretolamna appendiculata (Agassiz), upper lateral tooth in lingual view, PIL 85. G, H, Squalicorax kaupi (Agassiz), anterior tooth in labial (G) and lingual (H) views, PIL 71. I, J, Pseudocorax laevis Leriche, antero-lateral tooth in lingual (I) and labial (J) views, CLD 10. KeP, Squalicorax pristodontus (Agassiz). K, L, CLD 29, lateral tooth in labial (K) and lingual (L) views; M, CLD 47, lateral tooth in lingual view; N, O, CLD 4, anterolateral tooth in labial (N) and lingual (O) views; P, MSH 10, anterior tooth in lingual view. Scale bars represent 1 mm (A, B, D, E), 5 mm (C, F, I, J) and 10 mm (G, H, KeP).

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Maastrichtian of the USA, now has its geographical distribution extended to Europe. Order: Lamniformes Berg, 1958 Family: Anacoracidae Casier, 1947 Genus Squalicorax Whitley, 1939 Type species. Corax pristodontus Agassiz, 1843. Squalicorax kaupi (Agassiz, 1843) Fig. 3G, H For synonymy and bibliography, see Cappetta and Case (1975), Herman (1977) and Siverson (1992a), and add: 1843 Corax kaupi Agassiz, p. 225, pl. 26a, figs. 25e34; pl. 26, figs 4e8. 1977 Squalicorax kaupi (Agassiz); Herman, p. 107, pl. 4, fig. 2. 1989 Squalicorax kaupi (Agassiz); Mu¨ller, p. 43, fig. 10. 1989 Squalicorax kaupi (Agassiz); Williamson et al., p. 239, fig. 2f, g. 1992a Squalicorax kaupi (Agassiz); Siverson, p. 548, pl. 5, figs. 5e18. 1993 Squalicorax kaupi (Agassiz); Welton and Farish, p. 118, figs. 1e5. 1999 Squalicorax kaupi (Agassiz); Cappetta and Corral, p. 345, pl. 1, figs. 2, 3. 2000 Squalicorax kaupi (Agassiz); Bardet et al., p. 276, fig. 4m, o. 2001 Squalicorax kaupi (Agassiz); Cappetta and Odin, p. 647, pl. 1, fig. 1. 2001 Squalicorax kaupi (Agassiz); Gottfried et al., p. 494, fig. 2g. 2002 Squalicorax kaupi (Agassiz); Antunes and Cappetta, p. 119, pl. 7, figs. 9e12; pl. 8, figs. 4, 5. Material. 40 teeth: PIL 1e3, 15, 68, 70, 71, 81e84, 86, 90, 97, 100, 102, 105, CLD 1e3, 21, 28, 46, 57e59, MSH 31, 33, 34, 44. Numerous additional unnumbered, incomplete teeth. Localities and age. Le Pilou, Le Caillaud and Meschers; Middle Campanian (CIV and CV), Late Campanian (CVIII). Description. Serrated cutting edges, a strong labiolingual compression, a distinct distal heel and a gibbous mesial edge characterize these teeth, very often described and recognizable by their Galeocerdo-like morphology. They are relatively small with a mean width of about 10 mm. Anterior teeth (PIL 71; Fig. 3G, H) are higher than broad, unlike the lateral teeth, on which there is a slight reduction in the gibbosity of the mesial edge. A symphyseal tooth (PIL 11) has a triangular crown and

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no distal heel. The anaulacorhize root is relatively thick and approximately twice as high on the lingual side as on the labial side. The basal notch is shallow. Remarks. This well known cosmopolitan species constitutes a near-constant element of Campanian selachian faunas. A few of the largest teeth, still showing the characteristics of S. kaupi, correspond to intermediate forms located at the limit of S. kaupie S. pristodontus. Siverson (1992a, pl. 5, figs. 7, 8, 11, 12, 17, 18) figured similar teeth and also regarded them as being very close to S. pristodontus. Squalicorax pristodontus (Agassiz, 1843) Fig. 3KeP For synonymy and bibliography, see Cappetta and Case (1975) and Herman (1977), and add: 1843 Corax pristodontus Agassiz, p. 224, pl. 26, figs. 9e13. 1975 Squalicorax pristodontus (Agassiz); Cappetta and Case, p. 8, pl. 7, figs. 30e41. 1993 Squalicorax pristodontus (Agassiz); Welton and Farish, p. 119, figs. 1, 2. 1999 Squalicorax pristodontus (Agassiz); Cappetta and Corral, p. 345, pl. 1, fig. 1. 2000 Squalicorax pristodontus (Agassiz); Bardet et al., p. 276, fig. 4per. 2001 Squalicorax pristodontus (Agassiz); Gottfried et al., p. 494, fig. 2h. Material. Four teeth: CLD 4, 29, 47, MSH 10. Localities and age. Le Caillaud and Meschers; Middle Campanian (CV), Late Campanian (CVIII). Description. This species is distinguished from S. kaupi mainly by its large teeth and the attenuation of the distal heel, as in the specimen CLD 4 (Fig. 3N, O). Another tooth (MSH 10; Fig. 3P), from the Late Campanian of Meschers, displays a curve in lateral view (concave labial face) typical of certain teeth of this species (Welton and Farish, 1993, p. 119, fig. 1). The labio-lingual compression of the root is stronger than in S. kaupi. Remarks. The assignment of some large teeth from the Middle Campanian of Le Pilou (biozone CIV) to S. kaupi rather than to S. pristodontus remains for the moment more or less arbitrary. However, there is no doubt about the assignment of certain teeth from biozone CV and CVIII to S. pristodontus. The teeth of this species are definitely rarer in the Royannais than S. kaupi. Some teeth are questionably referred herein to S. pristodontus (CLD 29, 47; Fig. 3KeM), but could

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correspond to other undefined species. Only S. kaupi and S. pristodontus are considered in this study, pending a complete revision of the genus. Genus Pseudocorax Priem, 1897 Type species. Corax affinis Mu¨nster, in Agassiz, 1843. Pseudocorax laevis (Leriche, 1906) Fig. 3I, J 1906 Pseudocorax affinis var. laevis Leriche, p. 80 1977 Pseudocorax laevis Leriche; Herman, p. 114, pl. 4, fig. 4. Material. 12 teeth: PIL 35, 36, 91, 92, 96, 99, CLD 10, 11, 24, 25, 61e63. Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV and CV). Description. These medium-sized teeth (up to 11 mm) are very compressed labio-lingually and show sharp cutting edges devoid of serration. The anterior to antero-lateral teeth (CLD 10; Fig. 3I, J) show an erect cusp at the base of which a pair of lateral cusplets are found. The cusplets are rounded and largely fused to the main cusp. The lateral teeth approach more the morphology of Squalicorax; as the main cusp is inclined towards the rear, the mesial cusplet tends to disappear to form only one very slight nipple, and the distal cusplet forms a well-developed distal heel. The main cusp remains relatively narrow, which results in the teeth of Pseudocorax having a more gracile shape that those of Squalicorax. The root lobes are only well differentiated on the anterior teeth. A slight medio-lingual protuberance is present, from where a narrow but well-marked groove is observed. Remarks. This species is distinguished clearly from the Maastrichtian species P. affinis Agassiz by its lack of serration, and from the other CampanianeMaastrichtian species, P. granti Cappetta and Case, by its greater size and more robust morphology. P. laevis is well represented in the Campanian of the Anglo-FrancoBelgian Basin where it appears in the Turonian (Herman, 1977). In Europe, it also occurs in the Campanian of Sweden (Siverson, 1993b), and from the Coniacian to the Maastrichtian in North America (Hamm, 2001; Case and Cappetta, 2004). Pseudocorax also occurs in some Tethyan regions, such as Morocco and Israel (Arambourg, 1952; Lewy and Cappetta, 1989). Landemaine (1991) suggested that P. laevis originated from a population of Carcharias amonensis, with the resorption of cusplets and the formation of a distal heel leading to a convergence with Anacoraci-

dae, such as Squalicorax. The origin and systematic position of Pseudocorax requires a new analysis (Cappetta, pers. comm. 2004). More generally, the phylogenetic relationships of the different Anacoracidae remain very poorly understood. Family: Odontaspididae Mu¨ller and Henle, 1839 Genus Carcharias Rafinesque, 1810 Type species. Carcharias taurus Rafinesque, 1810. Carcharias latus (Davis, 1890) Fig. 4AeC 1890 Scapanorhynchus latus Davis, p. 386, pl. 38, figs. 14e17. 1890 Scapanorhynchus tenuis Davis (partim), pl. 38, fig. 12, non figs. 10, 11, 13. 1977 Scapanorhynchus? subulatus (Agassiz); Herman (partim), p. 185, pl. 7, fig. 5b, non fig. 5a. 1992a Carcharias latus (Davis); Siverson, p. 538, pl. 3, figs. 7e14. Material. 11 teeth: PIL 19e24, 73, 74, 79, 88, 101. Locality and age. Le Pilou and Le Caillaud; Middle Campanian (CIV). Description. These rather large teeth (up to 20 mm) have a relatively narrow and labio-lingually compressed crown. The lingual face is practically flat whereas the labial one is convex. The anterior teeth (PIL 20; Fig. 4C) have an erect main cusp with a pair of small, sharpedged cusplets that are inclined lingually. The lingual face is always completely smooth, while the labial one bears a very light basal ornamentation, made up of very short vertical folds, generally located below the cusplets. There is also a pinching of the enamel in the median part of the basilo-labial zone. The root has two distinct lobes, and the pronounced medio-lingual protuberance is divided by a distinct, deep groove. Lateral teeth (PIL 19; Fig. 4A, B) are broader and more flattened, and usually have a second pair of very small lateral cusplets. The cusplets are less sharp-edged than on the anterior teeth and widen to become triangular. The basilo-labial folds are very fine, but are clearer than on the anterior teeth, with a tendency to be present over the width of the crown. Unlike the lower lateral teeth, which remain relatively erect, the cusps of upper lateral teeth are rather clearly inclined towards the rear. Remarks. These teeth from the Middle Campanian of the Royannais correspond exactly to the descriptions given by Siverson (1992a) of C. latus from the Campanian of the Kristianstad Basin (Sweden), even if

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Fig. 4. AeC, Carcharias latus (Davis). A, B, PIL 19, upper lateral tooth in lingual (A) and labial (B) views; C, PIL 20, lower anterior tooth in lingual view. DeL, Carcharias adneti sp. nov. D, PIL 95, lower anterior tooth in lingual view; E, F, CLD 48, holotype, upper lateral tooth in lingual (E) and labial (F) views; G, H, PIL 50, lower antero-lateral tooth in lingual (G) and labial (H) views; I, J, CLD 17, lower anterior tooth in labial (I) and lingual (J) views; K, L, PIL 60, lower lateral tooth in lingual (K) and labial (L) views. MeP, Serratolamna khderii (Zalmout and Mustafa) comb. nov. M, N, MSH 3, upper antero-lateral tooth in labial (M) and lingual (N) views; O, P, MSH 1, lower lateral tooth in lingual (O) and labial (P) views. Q, R, Protolamna borodini (Cappetta and Case), antero-lateral tooth in lingual (Q) and labial (R) views, PIL 31. S, T, Cretolamna sp., upper lateral tooth, labial (S) and lingual (T) views, CLD 23. Scale bars represent 1 mm (IeL) and 5 mm (AeH, MeT).

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some Swedish specimens bear a few weak lingual folds. This species is also present in the Campanian of Belgium, illustrated by Herman (1977) under the name ?Scapanorhynchus subulatus Agassiz. Its discovery in south-west France extends its palaeogeographical distribution considerably southwards. Carcharias adneti sp. nov. Fig. 4DeL 2001 Carcharias sp.; Cappetta and Odin, p. 650, pl. 1, fig. 5. Material. 25 teeth: PIL 50e65, 80, 95, 106, 110, CLD 17, 18, 39, 40, 48. Numerous additional unnumbered, incomplete teeth. Derivation of name. Dedicated to Dr. Sylvain Adnet for his valuable help during the preparation of this paper, and for the washing and sieving. Types. Holotype, CLD 48; Paratypes, PIL 50, 60, 80, 95, CLD 17. Type locality and stratigraphic horizon. Middle Campanian (CIV) of Barzan, Le Pilou cliff (CharenteMaritime, France). Additional occurrences. Middle Campanian (CIV and CV) of Le Caillaud, Upper Campanian of Tercis (Landes, south-west France). Diagnosis. Species characterized by its small teeth (not exceeding 8 mm high) with a compressed crown with a flat labial face and a slightly convex lingual one. The main cusp is flanked by one pair of broad, relatively low, triangular lateral cusplets. A second pair of very reduced cusplets is often present on lower lateral teeth. The labial face is always completely smooth and there is a basal bulge slightly overhanging the root. The lingual face bears light and irregular vertical folds reaching the first third of the cusp. Folds are more or less marked and can be numerous (up to 40) according to the position of the tooth in the jaw, and completely missing in other teeth. Root lobes are rather short and with a flattened to slightly convex basal face. The basal margin of the root clearly shows an inverted V shape. Description. The holotype (CLD 48; Fig. 4E, F), which is rather broad and oblique, may correspond to an upper lateral tooth. It presents a compressed crown inclined towards the rear. The labial face is flat whereas the lingual one is slightly convex. The main cusp is flanked by a single pair of triangular marginal cusplets that are relatively large and weakly divergent. The labial face is completely smooth and a basal bulge clearly overhangs the root. The lingual face bears very light

short vertical folds. The root has a rather well-marked lingual protuberance with a deep groove, quite distinct lobes and a nearly flat basal face. The basal margin of the lobes is laterally rectilinear and medially angular. In anterior teeth (PIL 95, CLD 17; Fig. 4D, I, J), the main cusp and the cusplets are straighter and narrower. A small lower lateral tooth (PIL 60; Fig. 4K, L) has an additional minute outer pair of lateral cusplets. The root has a less marked lingual protuberance, less distinct lobes and a completely flat basal face. Remarks. Several species of Carcharias are known from Campanian and Maastrichtian rocks, notably in Europe and USA. Carcharias latus (Davis, 1890), C. gracilis (Davis, 1890) sensu Reynders (cf. Cappetta and Corral, 1999), C. steineri (Case, 1987) and C. sanguinei (Case, 1978) have more or less marked labial folds. Carcharias samhammeri (Cappetta and Case, 1975), C. heathi Case and Cappetta, 1997 and C. aasenensis Siverson, 1992a always have completely smooth labial and lingual faces. All of these species, therefore, can easily be distinguished from C. adneti sp. nov. Three species, C. hardingi, C. holmdelensis (Cappetta and Case, 1975) [ZC. cheathami (Case, 1987)] and C. tenuis (Davis, 1890 non Agassiz, 1843), resemble C. adneti with their ornamentation only being present on the lingual face. Teeth of Carcharias hardingi are larger, more robust, and have relatively smaller cusplets and more distinct root lobes. Teeth of C. holmdelensis have high, strong folds and more tapered cusplets, and those of C. tenuis have broad, triangular cusplets that are clearly attached to the main cusp. One tooth figured by Cappetta and Odin (2001) as Carcharias sp., from the Upper Campanian of Tercis (south-west France), corresponds perfectly to C. adneti and is herein assigned to it. Family: Serratolamnidae Landemaine, 1991 Genus Serratolamna Landemaine, 1991 Type species. Lamna serrata Agassiz, 1843. Serratolamna khderii (Zalmout and Mustafa, 2001) comb. nov. Fig. 4MeP 1952 Scapanorhynchus tenuis Davis; Arambourg, p. 52, pl. 4, figs. 1e20 2001 Carcharias khderii Zalmout and Mustafa, p. 388, pl. 1, figs. 1e3. 2001 Serratolamna serrata (Agassiz); Zalmout and Mustafa, p. 392, pl. 1, figs. 4, 5. 2002 Odontaspididae inde´t.; Antunes and Cappetta, p. 122, pl. 8, figs. 6e12. Material. 13 teeth: MSH 1e9, 12, 40e42.

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Locality and age. Meschers; Late Campanian (CVIII). Description. One of the best preserved specimens (MSH 3; Fig. 4M, N) corresponds to a medium-sized upper antero-lateral tooth (10 mm high). It has a triangular, narrow cusp, slightly tilted towards the rear and very compressed. Only the lingual face is slightly convex. A mesial cusplet and two distal ones flank this cusp. These cusplets are triangular, relatively broad and compressed labio-lingually. The mesial cusplet is erect and has a concave mesial edge. The first distal cusplet, like the main cusp, slightly bends towards the rear. The second distal cusplet is morphologically identical to the first, but very reduced. The labial face of the crown is completely smooth. The lingual face bears short vertical and a few marked folds, localised in the median part of the basal region. The root is not very thick and has a slight lingual protuberance carrying a distinct groove. A medio-labial depression is also present. The basal edge shows a broad notch. The basal face is relatively flat, especially at the end of the lobes. A lower lateral tooth (MSH 1; Fig. 4O, P) differs from the one described previously by its erect cusp and more marked basilo-lingual folds. The labial face of the crown displays a thick basal bulge that overhangs the root. The cusplets, two mesial and three distal, are divergent. Those of the first pair are definitely larger than the outer cusplets. The root has a more marked lingual protuberance and a deeper groove. The basal notch is definitely more concave and the lobes are flattened labio-lingually. Remarks. This species was described by Zalmout and Mustafa (2001) from the Lower Maastrichtian of Jordan and attributed to the genus Carcharias. They also figured a small lateral tooth they identified as Serratolamna serrata Agassiz. Examination of a large number of such teeth from the Lower Maastrichtian of Morocco (from where they were identified as Scapanorhynchus tenuis by Arambourg, 1952) and Jordan (H. Cappetta’s material) led me to conclude that they correspond to a unique taxon, Serratolamna khderii comb. nov. This differs from the type species of Serratolamna, S. serrata, by it smaller size and the presence of more or less marked lingual folds on most of the teeth. The diagnosis given by Landemaine (1991, p. 14) for the genus must therefore be corrected: ‘‘Cuspide ne pre´sentant pas d’ornementation, meˆme a` l’e´tat vestigial’’ [Crown devoid of ornamentation, even in a vestigial state]. This last character is not diagnostic and must be deleted. Fold-crowned S. khderii is the earliest known species of the genus, and may represent the ancestor of the larger, smooth-crowned, Maastrichtian S. serrata and S. caraibaea (Leriche). Specimens from the Upper Campanian of Meschers must be referred to this species, as well

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as the teeth recently figured by Antunes and Cappetta (2002) as indeterminate Odontaspididae from the Upper CampanianeLower Maastrichtian of Angola. In addition, the ‘‘small odontaspid’’ referred to Carcharias sp. by Bardet et al. (2000) from the Lower Maastrichtian of Syria also corresponds to this species (Cappetta, pers. comm. 2003). Family: Cretoxyrhinidae Glikman, 1958 Genus Cretolamna Glikman, 1958 Type species. Otodus appendiculatus Agassiz, 1843. Cretolamna appendiculata (Agassiz, 1843) Fig. 3F

For synonymy and bibliography, see Herman (1977) and Siverson (1992a), and add: 1843 Otodus appendiculatus Agassiz (partim), p. 270, pl. 32, figs. 2e8, 10e14, 16, 19, 20, 22e25, non figs. 1, 9, 15, 17, 18, 21. 1964 Lamna appendiculata (Agassiz); Albers and Weiler, p. 10, fig. 20. 1977 Cretolamna appendiculata (Agassiz); Herman, p. 210, pl. 9, figs. 2e4. 1992a Cretolamna appendiculata (Agassiz); Siverson, p. 528, pl. 1, figs. 1e17. 1993 Cretolamna appendiculata (Agassiz); Welton and Farish, p. 103, figs. 1e5. 1993 Cretolamna appendiculata (Agassiz); Williamson et al., p. 454, fig. 6.1e6. 1996 Cretolamna appendiculata (Agassiz); Siverson, p. 828, pl. 3, figs. 1e8. 1999 Cretolamna appendiculata (Agassiz); Cappetta and Case, p. 22, pl. 6, fig. 10, text-figs. 5, 6. 1999 Cretolamna appendiculata (Agassiz); Cappetta and Corral, p. 346, fig. 3. 2000 Cretolamna cf. appendiculata (Agassiz); Bardet et al., p. 276, fig. 4f, g. 2001 Cretolamna appendiculata (Agassiz); Gottfried et al., p. 495, fig. 2j. 2002 Cretolamna appendiculata (Agassiz); Antunes and Cappetta, p. 119, pl. 10, figs. 8e13. Material. 17 teeth: PIL 25e28, 69, 78, 85, 111e113, CLD 12e14, 41, 42, 44, 60. Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV and CV). Description. This species has relatively large teeth (up to 34 mm high). The anterior teeth have a triangular crown, which are rather broad mesio-distally and relatively compressed labio-lingually. The lingual face

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remains definitely more convex than the labial face, which is practically flat. Both faces are smooth. The cusplets are clearly detached from the main cusp, slightly divergent, always in a pair, broad and triangular. The root is massive and its lingual protuberance does not have any groove (secondary anaulacorhizie). The lobes are relatively distinct. The lateral teeth undergo a mesio-distal widening and a labio-lingual compression compared to the anterior teeth. The main cusp of the upper lateral teeth bends distally. The size of the cusplets increases (the mesial one is larger than the distal one) and become broader compared to the height. A lower lateral tooth (PIL 25) displays very short basal folds on its labial face that are rather localised below the cusplets. The root tends to take a quadrangular form, and in spite of an attenuation of its lingual protuberance, it remains thick and massive, with a convex basal face. An upper lateral tooth (PIL 85; Fig. 3F) has lateral cusplets with coarsely serrated cutting edges. Remarks. This cosmopolitan species is relatively uncommon in the Royannais, where it does not seem to reach the Upper Campanian. The teeth studied herein are very close, by their slightly divergent cusplets, to the populations attributed to the subspecies C. appendiculata pachyrhiza from the Campanian of the Obourg Chalk (Belgium) and the Kristianstad Basin (Sweden) (Herman, 1977; Siverson, 1992a). However, the use of subspecific taxa in palaeontology is unsatisfactory and not employed herein.

convex and concave respectively. The hypertrophy of the cusplets causes the latter to slightly overlap the labial face of the cusp. The root is nearly quadrangular in outline. A small notch on the basal edge and a light medio-labial depression however gives the impression of two very weakly-developed root lobes. The lingual protuberance does not project very much and the basal face is relatively planar, without a groove. Remarks. These teeth, unless they correspond to a teratological phenomenon, are clearly distinguished from C. appendiculata by their stronger labio-lingual compression, the narrowness of the main cusp, the hypertrophy, and the morphology of the cusplets. Similar cusplets are also present in C. nigeriana (previously regarded as being a subspecies, C. biauriculata nigeriana) from the Maastrichtian of Niger (Cappetta, 1972). This species is otherwise very close to the Royannais form. It differs mainly in the development of a second pair of cusplets. An incomplete tooth from the Upper Campanian of Tercis, south-west France (Cappetta and Odin, 2001), has distal cusplets that show the same morphological features as the two specimens from the Royannais, and is therefore tentatively referred to the same taxon. Genus Protolamna Cappetta, 1980 Type species. Protolamna sokolovi Cappetta, 1980. Protolamna borodini (Cappetta and Case, 1975) Fig. 4Q, R

Cretolamna sp. Fig. 4S, T ?2001 Cretolamna cf. appendiculata (Agassiz); Cappetta and Odin, p. 649, pl. 1, fig. 4. Material. Two teeth: PIL 77, CLD 23. Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV and CV). Description. The most characteristic tooth (CLD 23; Fig. 4S, T) is higher than broad. It presumably corresponds to an upper lateral tooth. Its cusp is relatively narrow, tilted towards the rear, and very compressed labio-lingually. The labial and lingual faces have the same low degree of convexity, and are completely smooth. The cusp is flanked by a pair of very well-developed divergent cusplets that reach a third of the height of the crown. They are also very compressed, triangular and entirely smooth. The apex of the mesial cusplet is rounded. The other cusplet, which is pointed and slightly smaller, is definitely tilted towards the rear. Its mesial and distal cutting edges are

For synonymy and bibliography, see Siverson (1992a) and add: 1975 Plicatolamna borodini Cappetta and Case, p. 23, pl. 3, figs. 1e9. 1992a Cretodus borodini (Cappetta and Case); Siverson, p. 534, pl. 2, figs. 21, 22. 2004 Protolamna borodini (Cappetta and Case); Case and Cappetta. Material. Ten teeth: PIL 31e34, 87, 94, 98, CLD 15, 16, 31. Localities and age. Le Pilou, Le Caillaud and Meschers; Middle Campanian (CIV and CV), Late Campanian (CVIII). Description. An antero-lateral tooth (PIL 31; Fig. 4Q, R), measuring 11 mm high, has a broad cusp at its base, tapering rapidly. The tooth is flanked by a pair of sharp-edged and very divergent cusplets. A second pair of tiny additional cusplets is developed.

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The whole of the crown has a slightly convex labial face and a very convex lingual face. The main cusp has concave edges. Its lingual face is completely smooth, which is not the case for the cusplets, which have slight vertical folds. The labial face has about 20 vertical, short, and very marked basal folds. These folds originate just above the base of the crown, and attenuate in the median part. The basilo-labial edge of the crown is very concave. The root is very high (more than half the total height of the tooth in lingual view). It has a very strong lingual protuberance, with no groove. The lobes are quite distinct and are subparallel and slightly compressed mesio-distally. The neck is clear. On the lateral teeth, as on the teeth of young individuals, some slight folds appear on the lingual face of the main cusp. Compared to the large anterior teeth, the labial folds go up higher, the cusplets are less divergent, and the root lobes are shorter. Remarks. This species, previously placed in Cretodus, displays all the characteristics of Protolamna (see Cappetta, 1980). The species ‘‘C.’’ borodini has been recently assigned to it (Case and Cappetta, 2004). The teeth of Protolamna occur in deposits ranging in age from Early Barremian (Kriwet, 1999) to Late Maastrichtian (Welton and Farish, 1993). Cappetta (1987b) and Lewy and Cappetta (1989) recorded a small species of Cretodus in the Upper Maastrichtian of Morocco and the Campanian of Israel respectively. It is also a species that belongs in Protolamna (Cappetta, pers. comm. 2003). In Europe, P. borodini is certainly present in the Campanian of Sweden (Siverson, 1992a). In an article dealing with a chimaeroid from the Upper Maastrichtian of Belgium, Duffin and Reynders (1995) noted C. cf. borodini within the selachian fauna. Now known from several species, Protolamna has long remained a monospecific genus because of the attribution of some of its species [e.g. P. compressidens (Herman)] to other genera. Together with the still-undescribed species, it appears that Protolamna was a significant element of Cretaceous selachian faunas. Family: Alopidae Bonaparte, 1838 Genus Paranomotodon Herman in Cappetta and Case, 1975 Type species. Oxyrhina angustidens Reuss, 1845. Paranomotodon sp. Fig. 5A, B 1845 Oxyrhina angustidens, Reuss, p. 6, pl. 3, figs. 7e 13. 1964 Isurus cf. angustidens (Reuss); Albers and Weiler, p. 9, fig. 33.

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?1975 Paranomotodon cf. angustidens (Reuss); Cappetta and Case, p. 24, pl. 5, figs. 1e10. 1977 Paranomotodon angustidens (Reuss); Herman, p. 189, pl. 7, fig. 7. 1992a Paranomotodon sp.; Siverson, p. 546, pl. 5, figs. 3, 4. Material. 18 teeth: PIL 37e49, 72, 89, 109, 114, CLD 43, MSH 11. Localities and age. Le Pilou, Le Caillaud and Meschers; Middle Campanian (CIV), Late Campanian (biozone CVIII). Description. The antero-lateral teeth, such as PIL 89 (Fig. 5A, B), are almost symmetrical and have an erect, rather narrow crown with a sigmoid profile. They can reach 17 mm in height. The lingual face is strongly convex and the two faces are completely smooth. On each side of the cusp the cutting edges are elongated to form short oblique heels, which are slightly lingually directed. The root has a strong lingual protuberance split by a deep groove. The root lobes are very developed and the basal face is strongly convex. The cusps of lateral teeth are more labio-lingually compressed and inclined towards the rear. The mesial edge is sigmoid and the distal one, above the heel, is practically rectilinear. The heels widen mesio-distally and are in the same plane as the cusp. Because of the slope of the tooth, the mesial heel tends to be based in the cusp while the distal heel becomes more distinct. The lingual protuberance of the root is more attenuated, the groove is less marked and the lobes are less distinct. Remarks. Siverson (1992a) figured a lower anterolateral tooth from the Campanian of Sweden. This is morphologically very close to specimens of the same tooth position from the Campanian of the Royannais. An unnamed species from the Lower Maastrichtian of New Jersey (Cappetta and Case, 1975) is characterized by its cusp being broader mesio-distally. Two teeth were also described from the Cenomanian of western Australia (Siverson, 1996). They differ from this material in their narrower cusp and more slender root lobes. The only species described, P. angustidens, is well known in the CenomanianeTuronian of Europe. As for the widespread Campanian forms of the genus, they have yet to be well defined. This can be explained by the low variability of the genus. Indeed, the Royannais teeth are not easily distinguishable as the P. angustidens illustrated by Herman (1977) from the Turonian, Coniacian and Santonian of Belgium. The assignment of Paranomotodon to the Alopiidae is retained herein, despite the opinion of Siverson (1996) who regarded it as incertae sedis.

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Fig. 5. A, B, Paranomotodon sp., antero-lateral tooth in lingual (A) and labial (B) views, PIL 89. CeF, Galeorhinus girardoti Herman. C, D, CLD 23, antero-lateral tooth in lingual (C) and labial (D) views; PIL 104, lateral tooth in labial (E) and lingual (F) views. G, H, Palaeogaleus faujasi (Geyn), lateral tooth in labial (G) and lingual (H) views, MSH 13. I, J, ‘‘Scyliorhinus’’ cf. germanicus Herman, lateral tooth in lingual (I) and labial (J) views, CLD 22. KeM, Scyliorhinus elongatus (Davis), antero-lateral tooth in lateral (K), lingual (L) and labial (M) views. Scale bars represent 1 mm (CeM) and 10 mm (A, B).

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Order: Carcharhiniformes Compagno, 1973 Family: Scyliorhinidae Gill, 1862 Genus Scyliorhinus Blainville, 1816 Type species. Squalus canicula Linnaeus, 1758. Scyliorhinus elongatus (Davis, 1887) Fig. 5KeM

For synonymy and bibliography, see Herman (1977) and add: 1887 Thyellina elongata Davis, p. 473, pl. 14, figs. 2, 3. 1977 Scyliorhinus elongatus (Davis); Herman (partim), p. 252, pl. 11, fig. 1d, e, g, i, j, non fig. 1aec, f, h. 1980 Scyliorhinus elongatus (Davis); Cappetta, p. 129, pl. 21; pl. 22; pl. 23, figs. 1e6, text-figs. 29e30. 1989 Scyliorhinus elongatus (Davis); Mu¨ller, p. 47, pl. 14, fig. 1. 1997 Scyliorhinus aff. elongatus (Davis); Noubhani and Cappetta, p. 58, pl. 24, figs. 1e6. Material. Two teeth: PIL 29, 115.

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less complete individuals (Davis, 1887; Cappetta, 1980), and in the Campanian of Belgium (Herman, 1977) and Germany (Mu¨ller, 1989). This species seems to reach the Upper Maastrichtian of Morocco (Noubhani and Cappetta, 1997). Hence, its rare occurrence in the Campanian of the Royannais is not surprising. ‘‘Scyliorhinus’’ cf. germanicus Herman, 1982 Fig. 5I, J

For synonymy and bibliography, see Mu¨ller (1989) and add: 1982 Scyliorhinus germanicus Herman, p. 141, pl. 2, fig. 10; pl. 4, figs. 4, 5. 1989 Scyliorhinus germanicus Herman; Mu¨ller, p. 47, pl. 11, fig. 5; pl. 12, figs. 1e4. 2001 Scyliorhinus sp.; Cappetta and Odin, p. 651, pl. 1, fig. 8. Material. One tooth deprived of its root: CLD 22. Locality and age. Le Caillaud; Middle Campanian (CV).

Locality and age. Le Pilou; Middle Campanian (CIV). Description. The single complete tooth (PIL 29; Fig. 5KeM) corresponds to an antero-lateral tooth, asymmetrical and distally inclined. It is 3 mm high and definitely higher than broad. The cusp is sub-conical, but with a lingual face that is more convex than the labial face. The first two-thirds of the labial face is ornamented by a score of vertical folds, rather fine but well marked. These folds go up higher along the edges than in the median area. The labial face has a strong medio-basal depression in which the crown overhangs the root. The lingual face is shorter because of the strong inclination of the tooth towards the interior of the mouth. It shows folds that are similar to those on the labial face. The cutting edges are difficult to distinguish among the folds. A pair of small, conical, divergent cusplets is situated at the base of the main cusp and largely fused to it. Both faces, from the base to the apex, are ornamented with fine folds. The root is rather thick and is broader than the crown. The lingual protuberance is very marked and split by a broad, deep groove. On the mesial side of this protuberance there are two large margino-lingual foramina (they also seem to be present on the distal side, although this is damaged). The basal face is flat. A strong medio-basal notch and a medio-labial depression delimit two asymmetrical lobes. The distal lobe displays a small extension directed towards the rear. Remarks. Scyliorhinus elongatus occurs in the Upper Santonian of Lebanon, where it is known from more or

Description. The single specimen recovered (CLD 22; Fig. 5I, J) corresponds to a small lateral, squat-crowned tooth, broader than high and slightly inclined towards the rear. In labial view, the sub-triangular crown shows concave cutting edges. On each side of the cusp, there are two pairs of cusplets. These are very reduced, far from prominent, and largely connected to the main cusp. The labial face is practically flat and slightly convex transversely at the level of the main cusp. The basal edge of this face is concave and forms a bulge overhanging the root (which is missing on this specimen). The labial ornamentation is not very marked and comprises some short, slightly flexuous and anastomosing medio-basal folds. In the marginal regions, these folds go up higher and are discontinuous. The lingual face is very convex, allowing the main cusp to be distinguished. In lateral view, this face is also slightly convex. It is more strongly ornamented than the other face. The marginal folds are more regular than the median folds. The absence of the root makes it possible to observe the basal crown face. The crown is trefoilshaped with one medio-lingual lobe corresponding to the main cusp, one to the mesial cusp and one to the distal cusp. Remarks. The general morphology of the single tooth is very close to some teeth of the type series from the Maastrichtian of Hemmor (Germany) (Herman, 1982, pl. 2, cf. fig. 10d). This species is also present in the Campanian of Germany (Mu¨ller, 1989) and in the

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Campanian and Maastrichtian of Sweden (Siverson, 1993b), and it seems to also be present in the Campanian of Belgium, having been illustrated by Herman (1977, pl. 11, fig. 1h) as Scyliorhinus elongatus. Cappetta and Odin (2001) figured an incomplete tooth from the Upper Campanian of Tercis, south-west France, as Scyliorhinus sp. This is morphologically identical to the Royannais specimen, and both certainly belong to the same species. The distribution of ‘‘S.’’ germanicus is thus limited to the Boreal area. If its presence in the Campanian of the Royannais is confirmed by the discovery of more complete specimens, it would be the most Tethyan occurrence. This scyliorhinid species does not belong to Scyliorhinus sensu stricto (Halter, 1995); it could be close to the Palaeogene genus Foumtizia Noubhani and Cappetta, or even belong to it. Indeed, the Royannais tooth suggests an affinity with certain teeth of Foumtizia gadaensis Noubhani and Cappetta (cf. Noubhani and Cappetta, 1997, pl. 32, fig. 3) because, for example, the basilo-lingual bulge is well developed and finely folded, the ornamentation is very reduced, and the cusplets are small. Family: Triakidae Gray, 1851 Genus Palaeogaleus Gurr, 1963

folds on the basilo-labial bulge. These folds are localised in the marginal zones of the bulge, under the mesial and distal heels. The root is thick and massive, with a weakly developed medio-lingual protuberance and lobes. The medio-lingual groove is relatively narrow and not very deep. The basal face is practically flat. Remarks. Palaeogaleus, known from Campaniane Ypresian deposits, has included, hitherto, four Cretaceous species: P. dahmanii Noubhani and Cappetta, Maastrichtian, Morocco; P. faujasi (van de Geyn), Maastrichtian, Belgium, Netherlands and Spain; P. havreensis Herman, Campanian, Belgium; and P. navarroensis Case and Cappetta, Maastrichtian, Texas. Of these, the Royannais tooth corresponds perfectly to the description of P. faujasi given by Herman (1977). P. havreensis, the only species of the genus known so far from the Campanian, is clearly distinguished from P. faujasi by its stronger and more isolated cusplets, and stronger basilo-labial folds, which are much more marked. The discovery of P. faujasi in the Upper Campanian broadens its stratigraphical distribution. Genus Galeorhinus Blainville, 1816

Type species. Scyllium vincenti Daimeries, 1888.

Type species. Squalus galeus Linnaeus, 1758.

Palaeogaleus faujasi (van de Geyn, 1937) Fig. 5G, H

Galeorhinus girardoti Herman, 1977 Fig. 5CeF

1937 Carcharhinus (Scoliodon) faujasi van de Geyn, p. 32, figs. 135e156. 1977 Palaeogaleus faujasi (van de Geyn); Herman, p. 261, pl. 12, fig. 2. 1999 Palaeogaleus faujasi (van de Geyn); Cappetta and Corral, p. 352, pl. 3, figs. 1e8.

1964 Synechodus cf. nierviensis Leriche; Albers and Weiler, p. 6, fig. 5aeb. 1977 Galeorhinus girardoti Herman, p. 268, pl. 12, fig. 7. 1997 Galeorhinus aff. girardoti Herman; Case and Cappetta, p.143, pl. 7, figs. 12, 13. Material. Two teeth: PIL 104, CLD 23.

Material. One tooth: MSH 13. Locality and age. Meschers; Late Campanian (CVIII). Description. The only tooth found (MSH 13; Fig. 5G, H) corresponds to a lateral tooth that is broader than high. The crown is broad and relatively massive. The main cusp, bent towards the rear, has a sigmoid mesial cutting edge and a rectilinear distal one. There is one pair of small rounded cusplets that are largely connected to the main cusp. The distal cusplet is more isolated than the mesial one. The marginal edges of the crown are also rounded. The labial face is very slightly convex in its median part. The basal edge is concave and forms a bulge that clearly overhangs the root. The lingual face is convex and completely smooth. The ornamentation of the crown is limited to the presence of very short vertical

Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV and CV). Description. The best specimen is a small anterolateral tooth (CLD 30; Fig. 5C, D) that is broader than high. It shows a cusp inclined toward the rear. The mesial cutting edge is slightly concave. There is a distal heel bearing three low cusplets, not well separated and decreasing in size distally. The labial face of the crown is flat whereas the lingual face has a strongly convex cusp. The labial and lingual faces are both smooth. A distinct basilo-labial bulge overhangs the root. Its enameloid is rather worn but seems to lack folds. The root is well developed and has a slightly rounded basal face. The lingual groove is broad and deep, with several foramina. In basal view, the mesial lobe is curved (U-shaped) while the distal one has a triangular outline. A lateral tooth

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(PIL 104; Fig. 5E, F) differs in having a more inclined cusp. Its distal heel displays a less distinct, single, cusplet. In addition, the basilo-labial bulge of the crown is less pronounced and smooth. The root is poorly preserved.

625

1977 Squatirhina kannensis Herman; Herman (partim), p. 157, pl. 6, fig. 10a, b, non fig. 10cej. 1987b Squatirhina lonzeensis Casier; Cappetta, p. 142, fig. 120dem. Material. Three teeth: PIL 30, CLD 27, 65.

Remarks. Although Galeorhinus occurs in various Late Cretaceous deposits (Cappetta, 1987b; Lewy and Cappetta, 1989; Siverson, 1993b; Welton and Farish, 1993), only two species have been described: G. girardoti from the CampanianeMaastrichtian of Belgium (Herman, 1977), Germany (Thies and Mu¨ller, 1993) and the USA (Case and Cappetta, 1997), and now the Campanian of western France, and G. glickmani from the Cenomanian of Russia (Popov and Lapkin, 2000). Superorder: Batomorphii Cappetta, 1980 Order: Rajiformes Berg, 1940 Suborder: Rhinobatoidei Fowler, 1941 Family: incertae sedis Genus Squatirhina Casier, 1947 Type species. Squatirhina lonzeensis Casier, 1947. Squatirhina lonzeensis Casier, 1947 Fig. 6AeC 1947 Squatirhina lonzeensis Casier, p. 13, pl. 5, fig. 2aed, text-fig. 4. 1977 Squatirhina lonzeensis Casier; Herman, p. 156, pl. 6, fig. 9.

Localities and age. Le Pilou and Le Caillaud; Middle Campanian (CIV and CV). Description. The only relatively well preserved specimen (PIL 30; Fig. 6AeC) is 3.5 mm wide and corresponds to a lateral tooth, being mesio-distally expanded and with a low cusp. In labial view, the cutting edges are concave. The labial face, moderately convex in its median part, is sub-rhombic and has a broad, not very strongly projecting apron, which is slightly bifid. The basal edge displays a strong bulge bearing some very short, vertical, marginal folds. Unlike the labial face, which is very tilted labio-lingually, the lingual face is erect and vertical. This has a broad sub-triangular median uvula, rounded at its end, and showing a strong concavity in lateral view. There are no lateral uvulae. This face also shows a distinct basal bulge where the very reduced ornamentation is similar to that observed labially. The cusp is worn and is very prominent. The holaulacorhize root is as wide as the crown. It has a flat, rhombic, basal face cut by a broad groove. A weak medio-lingual protuberance supports the uvula. The presence of margino-lingual foramina on both sides of this protuberance can be assumed.

Fig. 6. AeC, Squatirhina lonzeensis Casier, lateral tooth in occlusal (A), lateral (B) and basal (C) views, PIL 30. DeF, Rhinobatos echavei Cappetta and Corral, antero-lateral tooth in occlusal (D), lateral (E) and basal (F) views, PIL 93. Scale bars represent 1 mm.

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Remarks. Squatirhina occurs in AlbianeMaastrichtian deposits in northern Europe. Four species and an indeterminate form have been described: Squatirhina kannensis Herman, 1977, CampanianeMaastrichtian, Belgium and Holland; S. lonzeensis Casier, 1947, Turonian, Belgium and northern France (?), Santonian and Campanian, Belgium; S. thiesi Biddle, 1993, Albian, north-east France, Albian and Cenomanian of England; S. westfalica Mu¨ller and Dietrich, 1991, Cenomanian, Germany; Squatirhina sp., Campanian, Sweden (Siverson, 1993b). Some American and African forms have also been assigned to Squatirhina (Arambourg, 1952; Estes, 1964; Case, 1987), but they actually correspond to Orectolobiformes (Cappetta, 1987b). The study of a series of teeth of S. kannensis from the Maastrichtian of Maastricht (type locality) makes it possible to establish the major characteristics that clearly differentiate this species from S. lonzeensis. In S. kannensis, the labial face is flatter and the medio-lingual uvula is reduced (shorter and narrower). The margino-lingual foramina are larger in S. kannensis, and the basal folds are denser, finer, and can be developed at the level of the apron and the uvula. The Royannais tooth is very close to one of the teeth illustrated by Cappetta (1987b, fig. 120kem) and can thus be assigned to S. lonzeensis. This is also the case of two teeth illustrated by Herman as S. kannensis (1977, pl. 6, fig. 10a, b) from the Campanian Chalk of Obourg (Belgium). Indeed, the lateral tooth (fig. 10b) shows a long, broad uvula as well as a convex labial face. Moreover, the anterior tooth (fig. 10a) has distinct lateral expansions and does not show the strong mesio-distal compression of S. kannensis. S. lonzeensis thus reaches the Campanian, and this is the first record outside the Anglo-Franco-Belgian Basin. It is now well established that Squatirhina belongs to the Rhinobatoidei. It has even been placed in the family Rhinobatidae by some authors (Biddle, 1993; Underwood and Mitchell, 1999). Family: Rhinobatidae Mu¨ller and Henle, 1838 Genus Rhinobatos Linck, 1790 Type species. Raja rhinobatos Linnaeus, 1758 Rhinobatos echavei Cappetta and Corral, 1999 Fig. 6DeF 1999 Rhinobatos echavei Cappetta and Corral, p. 354, pl. 5, figs. 1e6. Material. Six teeth: PIL 76, 93, MSH 16, 32, 36, 37. Localities and age. Le Pilou and Meschers; Middle Campanian (CIV), Late Campanian (CVIII). Description. The Royannais teeth are rather massive and broader than long. The crown lacks a cusp and

shows a sharp transverse keel. In occlusal view, this is directed labially on both sides of the apex and thus shows a clear labial concavity. The labial face has a rhombic contour, with a convex basal edge. In lateral view, the visor is not very angular and its lower part displays a weak, rounded median process. One of teeth (MSH 16) shows some short, slight, vertical folds on each side of this process. The lingual face has a welldeveloped median uvula, rather broad and round at its end. The short, broad, rounded lateral uvulae are rather distinct. Just behind the marginal angles, there is a slight constriction of the lateral part of the crown. In lateral view, the labial face is clearly more convex than the lingual face. The root is only relatively well preserved on one of the six specimens (PIL 93; Fig. 6DeF). The root is rather thick and broad, and almost as high as the crown. The lobes, separated by a broad groove, have a relatively convex basal face. The groove contains a foramen in a rather labial position. A slight notch in the lingual contour of the lobes is present at the level of the single pair of margino-lingual foramina. The latter are well developed. Remarks. These teeth correspond perfectly to the description of Rhinobatos echavei from the Maastrichtian of the Spanish Basque Country (Cappetta and Corral, 1999). This species is distinguished from other Late Cretaceous Rhinobatos species by the combination of the following characters: teeth broader than long, lack of cusp, labial concavity of the transverse keel, presence of distinct lateral uvulae, and constriction behind the marginal angles. It is also very close to R. mariannae from the Upper Maastrichtian of the Netherlands and Belgium (Bor, 1983), but differs from it by its more developed and more rounded median and marginal uvulae. This new discovery of R. echavei, previously known only from the type locality, extends its stratigraphical range to the Campanian. It is possible that this species is among the many forms in the Maastrichtian of Morocco that are still awaiting study. Order: Myliobatiformes Compagno, 1973 Family: Rhombodontidae Cappetta, 1987b Genus Rhombodus Dames, 1881 Type species. Rhombodus binkhorsti Dames, 1881. Rhombodus carentonensis sp. nov. Fig. 7AeL

For an accurate description, the terminology used for this taxon is that employed by Noubhani and Cappetta (1994).

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Material. 31 teeth: PIL 107, CLD 19, 20, 32e36, 49e 53, MSH 17e30, 35, 38, 39, 43.

Types. Holotype, MSH 17; paratypes, PIL 107, MSH 35, 38, CLD 34, 35, 53.

Derivation of name. Latin, Carentonia, Charentes, after its geographical distribution in the Charentes.

Type locality, stratigraphic horizon and age. Upper Campanian (CVIII) of Meschers, Charente-Maritime,

Fig. 7. AeL, Rhombodus carentonensis sp. nov. AeC, MSH 35, antero-lateral tooth in labial (A), lateral (B) and basal (C) views; DeG, MSH 17, holotype, anterior tooth in labial (D), lingual (E), lateral (F) and occlusal (G) views; H, CLD 32, lateral tooth in lingual view; I, CLD 34, anterolateral tooth in labial view; JeL, CLD 35, lateral tooth, lingual (J), lateral (K) and basal (L) views. Scale bars represent 1 mm (H, JeL) and 5 mm (AeG, I).

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France; Middle Campanian (CV), Late Campanian (CVIII). Additional occurrences. Middle Campanian (CV) of Le Caillaud and Le Pilou, Upper Campanian (CVIII?) of Barbezieux, Charente, south-west France. Diagnosis. Relatively large species of Rhombodus characterized by its high-crowned anterior teeth. There is an angular crown with a plane occlusal face and sharp edges. The lingual bulge is slim and rectilinear. The ornamentation is identical on both labial and lingual faces and consists of distinct irregular vertical folds. The enameloid has fine small alveoli all over its surface. Description. The holotype corresponds (MSH 17; Fig. 7DeG) to an anterior or median tooth measuring 13 mm high and 9 mm wide. The crown, very slightly tilted lingually, is as high as it is wide. It is rhombic in occlusal view, square in labial and lingual views. The occlusal face is flat and very clearly delimited by sharp edges. These margino-labial and margino-lingual edges form an angle with the occlusal face slightly higher and lower than 90  respectively. The margino-labial faces are slightly concave and have rather irregular, very strong, sub-vertical folds in the lower half, becoming weaker towards the apex. The medio-labial edge is not very sharp and rectilinear in lateral view. The basilolabial transverse edge is prominent, almost rectilinear and cut out by the development of the many folds. It delimits the narrow, concave lower face of the visor. The margino-lingual faces are slightly convex and show the same type and frequency of folds and grooves. The medio-lingual edge is strongly projecting and convex. The lingual transverse depression is well marked. The lingual bulge is rectilinear, fine, and decorated with small vertical folds and bulges. The marginal edges extend to the base of the crown to form a small irregular triangular surface. The whole of the crown shows, for the unworn teeth, a very finely alveoli-ornamented enameloid. This forms a secondary ornamentation in addition to the labial and lingual folds. The root, lower and less broad than the crown, is rather high, not very thick and slightly lingually placed in lateral view. It has two triangular lobes separated by a broad, deep median groove. The state of preservation of the root does not allow observation of the presence of foramen. In lateral view, the labial face is oblique whereas the lingual face is vertical. The latter shows a clear neck or collar located under the basal bulge of the crown, as well as a light median protuberance overhanging the opening of the groove. Teeth from more lateral positions, such as specimens CLD 32 and 35 (Fig. 7H, JeL), are more or less rhombic and asymmetrical according to their position. They are more inclined mesio-distally and labio-

lingually but otherwise show the same characters as those of the anterior teeth. The occlusal face is always strongly worn, preventing the initial height of the crown from being observed. The root, always rather high and slightly more massive, has triangular lobes in basal view. The latter are sometimes trapezoidal owing to the truncation of the marginal parts. Four to five foramina open in the median area of the groove. The mean size of the teeth from the Middle Campanian is lower than that of the teeth from the Upper Campanian (about 6 and 9 mm respectively; maximum widths observed are 8 and 13 mm respectively). Remarks. The widespread Rhombodus was hitherto represented by five species restricted to the Maastrichtian: Rhombodus andriesi Noubhani and Cappetta, 1994, Upper Maastrichtian, Morocco and Spain; R. binkhorsti Dames, 1881, Maastrichian, cosmopolitan; R. levis Cappetta and Case, 1975, Lower Maastrichtian, New Jersey; R. meridionalis Arambourg, 1952, Lower Maastrichtian, Morocco, Egypt and possibly Syria; R. microdon Arambourg, 1952, Upper Maastrichtian, Morocco and Iraq. The teeth of Rhombodus carentonensis sp. nov. clearly differ from these other five species. R. levis has generally smaller teeth, with a smooth crown lacking a lingual bulge. The teeth of R. meridionalis are characterized by their more globular general aspect with a more definitely convex occlusal face, less projecting folds and less sharp edges. R. carentonensis shows more affinities with the more homogeneous group of R. binkhorsti, R. microdon and R. andriesi. It differs from the last two by its greater size, and from the largest species, R. binkhorsti, by its slimmer and rectilinear lingual bulge, its slighter ornamentation, and its unusual proportions. Indeed, the holotype does not enter into the morphometric variations of the Moroccan R. binkhorsti specimens (cf. Noubhani and Cappetta, 1994). The discovery of Rhombodus in the Campanian of the Royannais extends its stratigraphical distribution and marks its earliest occurrence. Indeed, it seems today that the stratigraphical position of North-American species R. levis is restricted to the Early Maastrichtian. MideLate Campanian specimens of R. levis from Georgia (Case and Schwimmer, 1988) appear, by their general morphology (relatively reduced root, high crown showing a sub-hexagonal contour) and their very small size (2.5 mm broad), to be close to the rhinobatoid genus Hypsobatis Cappetta. The occurrence of a true species of Rhombodus in the Campanian may have biostratigraphical implications in numerous regions, because this genus had been previously unknown worldwide in deposits older than Maastrichtian and was therefore thought to be characteristic of this stage (Bardet et al., 2000). In spite of their Campanian age, the teeth of R. carentonensis are well developed and very characteristic.

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The presence of essentially modern rhombodontid teeth in the Campanian suggests that the Rhombodontidae is a more ancient group than had previously been recognized. Interestingly, the grinding family Myliobatidae seem also to have appeared during the Early Campanian in North America, with Brachyrhizodus (Cappetta and Case, 1975; Cappetta, 1987a,b; Welton and Farish, 1993) and an indeterminate Texan form close to Myliobatis (Welton and Farish, 1993, p. 154, figs. 1e2). Differentiation, within Myliobatiformes, of taxa with grinding type teeth (Myliobatoidea: Rhombodontidae and Myliobatidae) thus seems synchronous with the Campanian diversification of Rhinobatoidei of crushing-grinding rank observed among Hypsobatidae (Cappetta, 1992). The decline and Campanian extinction of the Ptychodontidae (Cappetta, 1987a), hybodontoid sharks with a very specialized grinding type dentition (Macleod, 1982; Williamson et al., 1991), was surely one of the principal factors, having involved simultaneously and by convergence, this adaptive tendency relating to the teeth of many Rajiformes and Myliobatiformes. R. carentonensis also occurs in the Upper Campanian (CVIII?) of Barbezieux, in Charente (pers. obs.).

4. Conclusions The elasmobranch species recovered from the upper half part of the type Campanian are summarized in

CIV

Fig. 8 and their stratigraphical distributions indicated. Even if some species (mainly the smallest ones) are represented by very reduced numbers of specimens, which leads to a biased sample, a perceptible faunal change can be observed between the Campanian CIV and CVIII. On the one hand, this evolution of the selachian fauna is characterized by rather numerous local extinctions at the end of biozone CIV time (three extinctions) and more especially during the CVIeCVII gap (height local extinctions). On the other hand, Rhombodus carentonensis sp. nov. clearly occurs only just above the CIV/CV boundary, and two species, Serratolamna khderii comb. nov. and Palaeogaleus faujasi, seem to have appeared in the region during the Late Campanian. Lastly, Paranomotodon sp. and Rhinobatos echavei seem to be absent from the Campanian CV and thus prove to be Lazarus taxa. However, concerning the small species known only by a very few specimens (e.g., Scyliorhinus elongatus and P. faujasi), these observations may result from a collecting bias, as mentioned above. The deepening trend recorded in biozone CV may certainly have governed the evolution of the selachian fauna. Indeed, this environmental change may have produced definitive or temporally local disappearances within benthic/necto-benthic species rather restricted to the inner neritic habitats, accompanied by a development of large pelagic forms (not only sharks, but also actinopterygians and mosasaurids, pers. obs.). Besides, such a pattern of faunal changes has been previously described among irregular

CV

CVI CVII

CVIII

Plicatoscyllium cf. globidens Carcharias latus Scyliorhinus elongatus Squatina cf. hassei Pseudocorax laevis Carcharias adneti sp. nov. Cretolamna appendiculata Cretolamna sp. Galeorhinus girardoti Squatirhina lonzeensis Heterodontus cf. rugosus Squalicorax kaupi Protolamna borodini Paranomotodon sp. Rhinobatos echavei Squalicorax pristodontus "Scyliorhinus" cf. germanicus Rhombodus carentonensis sp. nov. Serratolamna khderii Palaeogaleus faujasi

Fig. 8. Stratigraphic occurrences (biozone level) of type Campanian selachians within the part of the succession studied (biozones CIVeCVIII, except for biozones CVI and CVII).

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echinoids and asteroids of the type Campanian (Ne´raudeau and Villier, 1997; Villier et al., 1997) and more widely in the Campanian of the Tethyan realm within rudists (Philip, 1985). A hypothetical warming of the marine waters, possibly correlated to this glacio-eustatic transgression, could explain the extinction of the Boreal species Carcharias latus. Moreover, it could also have favoured the emigration from tropical waters (Texas or southern Tethys) of a crushing or grinding-feeding myliobatiforme stock which then might have been at the origin of R. carentonensis. Palaeogeographically, the Charentes region is intermediate between the Boreal and Tethyan realms, as is the Maastrichtian Albaina locality in the BasqueCantabrian Basin (Cappetta and Corral, 1999). Unfortunately, and contrary to the Maastrichtian, the Campanian selachian faunas are still poorly known in Southern Europe and North Africa (Cappetta, 1989). Thus, a comparison in order to understand and establish the modalities of selachian fauna migrations remains difficult for the Campanian. However, it is notable that the Charentes fauna was dominated during the Middle Campanian by an important community of large lamniforms, well represented in Northern Europe (Herman, 1977; Siverson, 1992b), and associated with some other Boreal taxa, such as Squatirhina. During the Late Campanian, an inversion of this situation took place, with the fauna showing numerous Tethyan affinities (e.g., Serratolamna khderii). This change is surely in relation with the contemporaneous closure of the BasseLoire Strait which isolated the Aquitain Basin from the Paris Basin and then directly stopped northern immigrations. It is also interesting to observe that six species out of eight recovered from Meschers are identical or closely related to those described from the Maastrichtian of Albaina. Indeed, it is reasonable to suppose that the development of Rhombodus carentonensis and the appearance of S. khderii added to the early occurrence of Palaeogaleus faujasi and Rhinobatos echavei in the Aquitano-Pyrenean region (palaeogeographical unit sensu Philip, 1985) constitute elements related to the origin of the Maastrichtian Basque-Cantabrian fauna. During the Late Cretaceous, squaloid and palaeospinacid sharks were particularly well represented in the cooler and/or deeper waters of North European basins (Germany, Sweden) (Herman, 1982; Mu¨ller and Scho¨llmann, 1989; Mu¨ller, 1989, 1991; Siverson, 1992b, 1993a; Thies and Mu¨ller, 1993), as in the northern part of the Western Interior Seaway (North America) (Case, 1978, 1987). Their absence from the Campanian of Charentes is therefore not surprising according to the environmental and climatic conditions then existing in the Aquitaine Basin, as mentioned above. By contrast, the absence of sclerorhynchid sawfish, usually present in most of Late Cretaceous selachian faunas, is significant and rather unexpected. Indeed, this group occurs in the

Santonian of Vende´e (Cappetta, 1981) and Maastrichtian of the Basque-Cantabrian region (Cappetta and Corral, 1999), two areas close to the Charentes region. As for the hybodontoid sharks, also absent from the Royannais fauna, their last representatives in the Aquitaine Basin seem to occur in the Late Santonian paralic deposits of Dordogne (Landemaine, 1991), with Polyacrodus sp. Lastly, it is interesting to note the absence of the cosmopolitan Late Cretaceous genus Scapanorhynchus from the Middle and Upper Campanian biozones dealt with herein, even though it occurs in Lower Campanian biozone CIII in Charentes, based on the discovery of a single tooth from a small outcrop near Barzan (pers. obs.). In conclusion, this study has improved our knowledge of the Campanian elasmobranch fauna in the periTethyan regions, and more particularly in the stratotype area in the northern Aquitaine Basin. Among the 20 species recognized, two are new, and the geographical and stratigraphical ranges of numerous taxa are now extended. This is especially important with respect to the world-wide genus Rhombodus, because its first discovery in Campanian layers indicates that it is not confined Maastrichtian rocks, as previously thought.

Acknowledgements I am particularly grateful to everyone who made possible the long field work and the collecting of the specimens discussed herein, and especially my parents, R. and D. Vullo, for their valuable assistance and patience. I am most grateful to all those who put at my disposal or gave me their material, especially P. and A. Be´ne´fice, T. Lenglet, J. Mullon and E. Ruhaud. S. Adnet and H. Cappetta, University of Montpellier, and P. Miramand, University of La Rochelle, are thanked for various kinds of assistance (documentation, photographic and SEM work) suggestions and discussions. I also thank D. Ne´raudeau, University of Rennes, and L. Villier, University of Marseille, for their valuable observations on the stratigraphy of the type Campanian, as well as N. Bardet (Muse´um National d’Histoire Naturelle, Paris) for the determination of the mosasaur material. Lastly I thank the two reviewers, H. Cappetta and C.J. Underwood, for their helpful comments, and M. Hill, for her help with the English version.

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