An Early Campanian vertebrate fauna from the Villeveyrac Basin (Herault, Southern France)

An Early Campanian vertebrate fauna from the Villeveyrac Basin (Herault, Southern France) By Eric Buffetaut, Paris, Guy Costa, Montpellier, Jean Le Loeuff, Esperaza, Michel Martin, Boulogne-sur-Mer, Jean-Claude Rage, Paris, Xavier Valentin, Vitrolles, and Haiyan Tong, Paris BUFFETAUT,E.; COSTA,G.; LE LOEUFF,].; MARTIN,M.; RAGE,].-C.; VALENTIN,X. & TONG,H. (1996): An Early Campanian vertebrate fauna from the Villeveyrac Basin (Herault, Southern France). - N.]b. Geol. PaUiont. Mh., 1996 (1): 1-16; Stuttgart. Abstract: A rich vertebrate fauna from the Campanian of Villeveyrac includes fishes (Sparidae, Lepisosteidae), anurans (with the oldest known representative of the Palaeobatrachidae), chelonians (Podocnemidinae), lizards, crocodiles (Trematochampsidae, indeterminate Eusuchia) and dinosaurs (Theropoda, Ornithopoda, Ankylosauria). The discovery of this assemblage leads to some paleoecological a!1d biostratigraphical hypotheses. The absence of titanosaurid sauropods in the Early Campanian European faunas is discussed. Zusammenfassung: Eine reiche Wirbeltierfauna aus dem Campan von Villeveyrac enthiilt Fische (Sparidae, Lepisosteidae), Frosche (der iilteste bekannte Vertreter der Palaeobatrachidae), Schildkroten (Podocnemidinae), Eidechsen, Krokodilier (Trematochampsidae, unbestimmte Euschier) sowie Dinosaurier (Theropoda, Ornithopoda, Ankylosauria) und erlaubt einige palokologische und biostratigraphische Annahmen. Die Abwesenheit von Titanosauridae im Untercampan Europas wird diskutiert. Resume: La description d'une riche faune de vertebres du Campanien inferieur de Villeveyrac, comprenant des poissons (Sparidae, Lepisosteidae), des amphibiens (Ie plus ancien Palaeobatrachidae connu), des tortues (Podocnemidinae), des lezards, des crocodiles (Trematochampsidae, Eusuchia indetermines) et des dinosaures (Theropoda, Ornithopoda, Ankylosauria) permet quelques hypotheses paleoecologiques et biostratigraphiques. L'absence des Titanosauridae dans les gisements du Campanien inferieur d'europe est commentee. Introduction The Late Cretaceous vertebrates from southern Europe have long been neglected by paleontologists. A few years ago, our team started both a systematic revision of old collections and systematic excavations at 1 N. Jb. GeoL Paliionl. Mh. 1996 0028-3630/96/1996-0001 $ 4.00 1996 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart

several localities in southern France and northern Spain (LE LOEUFF 1992). At the time of writing, the best known localities are in Maastrichtian deposits in southern France (BUFFETAUTet al. 1989), the Spanish Basque Country (ASTIBIA et al. 1990) and Transylvania (WEISHAMPELet al. 1991). A few Early Campanian continental vertebrates have been reported from Sweden, Austria and southern France. In Sweden, the Blacksudden locality (Scania) has yielded a single theropod tooth (PERSSON1959); an ornithopod tooth has been briefly reported from the Lower Campanian of Scania by NEssov (1993). In Austria, the Muthmannsdorf locality has yielded a diverse fauna (BUNZEL 1871, SEELEY 1881, NOPcsA 1926), including remains of the nodosaurid ankylosaur Struthiosaurus austriacus (PEREDA-SUBERBIOLA& GALTON1992), a small ornithopod dinosaur (Rhabdodon cf. priscus), a possible hsisosuchid crocodile (Doratodon carcharidens) and an alligatorid (BUFFETAUT 1979), a pterosaur (WELLNHOFER1980) and a champsosaur (BUFFETAUT 1989a). This locality (a lignite mine) is no longer accessible. In Provence, a few "Fuvelian" (i. e., Early Campanian) localities (see BUFFETAUT& LE LOEUFF1991, for a discussion of the age of the Late Cretaceous continental deposits of Europe) have yielded vertebrate remains. They are essentially chelonian, crocodilian and fish remains (MATHERON 1869, LE LOEUFF1992). A new abelisaurid theropod (Tarascosaurus salluvicus) has recently been described from the Campanian of Le Beausset (LE LOEUFF& BUFFETAUT1991). The newly discovered vertebrates from Villeveyrac constitute the best known Early Campanian assemblage from Western Europe. Geological setting The Villeveyrac basin is situated between Sete and Pezenas in the "departement" of Herault (Fig. 1). It is about 20 km long from South West to North East, and 5 to 6 km wide. Structurally it comprises the Castelnau de Guers anticline and the Villeveyrac syncline. The Late Cretaceous continental deposits (FREYTET 1970, 1971) are situated above an Albo-Cenomanian bauxite (BERGERet al. 1981). The age of those deposits has long been disputed. According to FREYTET(1970, 1971), the lower, lignite-bearing grey clays (which have yielded the vertebrate remains described in the present paper) contain an invertebrate fauna which permits a correlation with the Fuvelian of Provence, whereas the upper, more sandy, part would be an equivalent of the "Begudo-Rognacian" (i. e., Maastrichtian), also on the basis of its invertebrate fauna. According to FEIST& FREYTET(1983), pollen, ostracodes and charophytes all indicate a "Fuvelian" age for the lower part of the series. The palaeoecology of the basal series of the Late Cretaceous beds of the Villeveyrac

~:-"'----' [ --,_1 Fig.!. Location map showing the department of Herault in southern France and the location of Villeveyrac in Herault. basin has been studied in detail by FREYTET(1970, 1971), who reconstructed an estuarine landscape combining freshwater and brackish environments. Fossil vertebrates were reported from the Upper Cretaceous of the Villeveyrac basin as early as 1872 by BLEICHER,who mentioned the occurrence of "large saurians" and "iguanians?" in sandstones and marls he considered as an equivalent of MATHERON'S"Etage de Rognac" in Provence. BLEICHER'Smaterial was seen by GERVAIS,who identified some vertebrae as probably belonging to a dinosaur, and considered that the fauna was comparable with that from MATHERON'S"Etage de Fuveau" (GERVAIS1872). GERVAISlater referred the vertebrae from Villeveyrac to Rhabdodon (GERVAIS 1877). In 1947, LAPPARENTmentioned" scattered bones from the red sandstones in the upper part of the Late Cretaceous series. Where the older finds came from is not perfectly clear, but it seems that most of the specimens reported in the nineteenth century (and now lost) may have come from the same levels as those mentioned by LAPPARENT, i. e. from the Maastrichtian. To the contrary, the vertebrate remains found in recent years in the Villeveyrac basin come from the dark clays and associated more sandy facies of the lower, Campanian, levels. The washing and screening of sediment from the Villeveyrac basin resulted in the description of a single theropod tooth a few years ago (BUFFETAUTet al. 1986). Renewed collecting in recent years has yielded an important new material which is described below. Most of the described specimens or casts of them have been deposited in the collections of the Musee des Dinosaures in Esperaza (Aude).

(irb Fig. 2. Fish remains from the Villeveyrac Basin. A-D: Lepisosteid (Early Campanian, Villeveyrac). A: tooth (MDE-Vii-03), scale bar: 5 mm; B: tooth (MDE-Vil-04); C: apex of the tooth MDE ViI-OS; D: basis of MDE-Vil-03, scale bars: 2 mm. E-F: Teeth of Lepisosteus (Late Cretaceous, Portugal) redrawn from SAUVAGE (1897-98); scale bar: 10 mm. G-K: Scales of lepisosteids (Early Campanian, Villeveyrac). G: MDE-Vil-18; H: MDE-Vil-07; I: MDE-Vil-08; J: MDE-Vil-19; K: MDE-Vil-20; scale bars: 10 mm. L-N: Scales of Lepisosteus (Late Cretaceous, Portugal) redrawn from SAUVAGE (1897-98); scale bars: 10 mm. a: Scale oflepisosteid (Early Campanian, Villeveyrac) MDE-Vil-06; scale bar: 10 mm. P: Part of the lower jaw of living Sparus aurata (LINNE, 1758); scale bar: 10 mm. Q:-R: Teeth of Sparidae indet. (Early Campanian, Villeveyrac). Q Basal face of MDE-Vil-09, scale bar: 2 mm; Ra: apical face ofmde-vil-10; Rb: side view of MDE-Vil-10; scale bar: 2 mm.

Faunal list Teleostei Sparidae indet. (Fig. 2P, Q R) Several tiny button-like teeth (Fig. 2) have been obtained by washing and screening sediment. This type of tooth morphology is recorded in different groups of fishes but on the basis of radiating grooves (Fig. 2P, Q) ornamenting the basal face of the teeth they are referred to the family Sparidae. The living Sparus aurata usually inhabits shallow marine near-shore environments but this species (and other related ones) commonly also lives in brackish water..., Ginglymodi Lepisosteidae indet. (Fig. la-o) Some fish remains from Villeveyrac are referred to a lepisosteid. They include some incomplete large teeth (Fig. 2A) exhibiting "plicidentine" (Fig. 2D) and complete (Fig. 2B) or fragmentary (Fig. 2C) smaller lanceolate teeth. Although they are very similar to those of Lepidotes, scales from different parts of the body (Fig. 2E, F) are referred to a lepisosteid since the locality has yielded no Lepidotes teeth. The living genus Lepisosteus is a freshwater fish and does not live in salt water. It is now restricted to North and Central America and Cuba. Fossil remains are recorded from Europe, Africa, South America and India (GAYET1987). Fragmentary lepisosteid remains were des.cribed from the Upper Cretaceous of Portugal as Clastes lusitanicus and C. pustulosus by SAUVAGE(1897-98). Clastes is now regarded as a junior synonym of Lepisosteus or Atractosteus, depending on the species considered (WILEY 1976). The generic status (Lepisosteus or Atractosteus) of the Portuguese material is disputed (see WILEY 1976, and GAYET1987, for conflicting opinions). In any case, on the basis of the fragmentary remains available no clear systematic or phylogenetic relationships between SAUVAGE'S species (if valid) and the lepisosteid from Villeveyrac can be ascertained. The Villeveyrac material is considered here as indeterminate at the generic and specific levels. Anura The locality has yielded only two remains belonging to anuran amphibians: An indeterminable fragment of tibiofibula (COSTAcollection, cast MDE-ViI-15) and a fragmentary frontoparietal (Fig. 3) which is referred to the Palaeobatrachidae.

Palaeobatrachidae indet. This frontoparietal (COSTA collection, cast MDE-Vil-16; Fig. 3) is represented by a (? posterior) half. It is azygous, elongate and thick. The mid part is narrow and the bone widens toward the extremity. The ventral face is rather concave. Traces of endocranial pattern and parietal foramen are hardly visible. There is apparently no pit for the pineal organ. Pineal and parapineal tracts (SPINAR1976) are conspicuous in the sagittal plane. The dorsal surface is flat and sculptured. On each side it is bordered by a narrow, smooth and concave area which is sloping latero-ventrally. The sculpture comprises few well-defined tubercules on the two latero-? posterior areas; the remaining part of the dorsal face shows circular traces and irregular pits. It may be questioned whether these traces and pits correspond to abraded tubercles. Ventrally, the latero-? posterior parts display a ridge and groove system. The Palaeobatrachidae are the only extinct family of anurans. Excepting perhaps the latest forms (Plio-Pleistocene) they were aquatic frogs living in fresh water. It is generally assumed that palaeobatrachids are known from the Late Jurassic (or Early Cretaceous) to the middle Pleistocene. But the allocation of the oldest fossils is questionable. Eobatrachus agilis, from the Late Jurassic of Wyoming, is known only by a humerus; its systematic position remains non-established (ESTES & REIG 1973). Montsechobatrachus gaudryi, from the Late Jurassic or early Cretaceous of Spain, is represented by a poorly preserved skeleton of difficult study. Although Fig. 3. Palaeobatrachidae indet. (Early Campanian, Villeveyrac); frontoparietal in dorsal (A) and ventral (B) views; scale bar: 5 mm.

originally assigned to the genus Palaeobatrachus, any precise allocation of this fossil within anurans would be dubious (HECHT 1963). Neusibatrachus wiiferti, another frog from the Late Jurassic or Early Cretaceous of Spain, has been considered a palaeobatrachid (ESTES& REIG 1973, SPINAR1975). However, it does not show any of the derived characters of the family. It could be regarded close to the ancestry of palaeobatrachids but the assignment to the family cannot be demonstrated. Furthermore, in a brief report, SANCHlZ& ROCEK(1992) have indicated that the oldest Palaeobatrachidae come from the Late Cretaceous which implicitly excludes these fossils from the family. Before the discovery of the Villeveyrac fauna, two Palaeobatrachidae only were known from the Late Cretaceous: Palaeobatrachus occidentalis from the late Maastrichtian of Wyoming and Montana (ESTES& SANCHIZ 1982), the only non-european representative of the family, and an indeterminate palaeobatrachid, of apparently early Maastrichtian age, from Spain (ASTIBIAet al. 1990). Consequently, if the exclusion from the family of the three Late Jurassic and/or Early Cretaceous fossils is right, then the specimen from Villeveyrac is the oldest representative of the Palaeoba trachidae. Chelonia Pleurodira Pelomedusidae Podocnemidinae Polysternon provinciale (MATHERON1869) An almost complete shell, with both carapace and plastron (COSTA Collection, Fig. 4), and many isolated plates are known from Villeveyrac (a locality where chelonian remains are especially abundant). The shell is about 50 em long, low and oval in shape, decorated with thin more or less dichotomised vascular grooves following the pelumedusoid pattern sensu BROIN (1977). The caracters of this specimen agree with those of Polysternon provinciale (MATHERON1869), from the Campanian of the Fuveau Basin (PORTIS1882, NOPCSA1931, BROIN 1977), with only a few minor differences. The size of our specimen is slightly larger than that of the shell described by NOPCSA(1931), the carapace seems to be wider. Vertebrals 2 to 4 are slightly broader than long, whereas they are longer than broad in NOPCSA'S specimen. On the plastron, the" anterior lobe appears to be clearly shorter than the posterior one, instead of only slightly so in NOPCSA'Sspecimen. The entoplastron is large, but its length is only 1,41 times as long as the distance between the entoplastron and

Fig. 4. Polysternon provinciale (Early Campanian, Villeveyrae); shell in ventral (A) and dorsal (B) views; scale bar: 10 em. the pectoro-abdominal sulcus, instead of twice as long as described by BROIN (1977) for NOPCSA'S specimen. The anal notch is more or less diamond-shaped, deeper and narrower than in the above-mentioned specimen, in which it is semi-circular in shape. The intergular is large, largely enters the entoplastron and widely separates the gulars, which are narrower than the intergular. There are no additional small gulars and the gulars do not reach the entoplastron. In NOPCSA'Sspecimen, the intergular is narrower than the gulars and hardly enters the entoplastron; posteriorly, the gulars reach the entoplastron. All these differences may be the result of individual variation and do not justify the erection of a new species. An almost complete skull (Fig. 5) has also been found at Villeveyrac (COSTA collection, cast MDE-Vil-12). It is about 8 cm long, with moderately large dorsal posterior notches. The well developed trochleal processes of the pterygoids and the presence of a medial process on the quadrates show that it belongs to a pleurodiran. The absence of the nasals and the prefrontals meeting on the midline allow to refer this skull to the Pelomedusidae. The enlarged ventral opening of the internal carotidian canal indicates affinities with the family Podocnemidinae. This skull very probably belongs to Polysternon provinciale, a form in which the skull was hitherto unknown. It will be described in detail in a separate paper (TONG, in prep.).

Fig. 5. Polyslernon provinciale (Early Campanian, Villeveyrae); skull in dorsal (A) and ventral (B) views; scale bar: 2 em. Cryptodira Dermatemydidae Many isolated turtle plates from Villeveyrac exhibit a marked vermiculate ornamentation. This ornamentation is reminiscent of some representatives of the Dermatemydidae, such as Tretosternon OWEN 1842, from the Purbeck and Wealden of England. Chelonians with a similar ornamentation have been reported from many Late Cretaceous (Campanian and Maastrichtian) localities of southern France' and northern Spain. Some of them have been referred to Apholidemys gaudryi MATHERoN1869 (see, for instance, BABINOTet al. 1983), a form originally described from "the base of the Rognac stage" of Provence by MATHERoN (1869). The shell of this turtle was described by MATHERONas "com me sillonnee en long par des rugosites irregulieres et tres saillantes" ("longitudinally furrowed by irregular and very prominent rugosities"). However, attribution to the genus Apholidemys, originally described by POMEL(1847) from the Cuisian of Cuise- La- Motte, is very uncertain, all the more so that POMEL'Soriginal material is lost and that, according to BRaIN (1977), the genus must be redefined. MATHERON'smaterial was considered by BROIN(1977) as probably belonging to a new cryptodiran genus of uncertain affinities at the family level. Plates of this type from Lano in the Spanish Basque Country have been referred to dermatemydids (ASTIBIAet al. 1990), an attribution considered as uncertain by LE LOEuFF (1992). Unfortunately, with the exception of some plates from

Montseret (Upper Cretaceous of the Corbieres; see TONG et al. 1993), none of these specimens have yet been figured. The available material from Villeveyrac is too fragmentary to warrant a precise identification. Squamata Lacertilia indet. A fragment of bone bearing two pleurodont teeth (MDE-Vil-11) evidences the presence of Lacertilia. It does not permit an assignment within the group. Crocodylia Mesosuchia Trematochampsidae indet. Trematochampsids are represented by stout blunt teeth with an enamel ornamented with sinuous ridges and granulous carinae. These teeth are completely similar to those from the Fuvelian of the Fuveau basin, which were named Crocodylus affuvelensis by MATHERON(1869), and in fact probably belong to a trematochampsid mesosuchian (BUFFETAUT 1989b), for which a new generic name will be proposed elsewhere. Eusuchia indet. A few procoelous vertebrae (COSTAcollection) indicate the presence of an indeterminate eusuchian crocodile in the Villeveyrac fauna. Very poorly known eusuchians are also present in,the "Fuvelian" lignites of Provence (BUFFETAUT1980). Dinosauria Theropoda indet. Several serrated and laterally compressed teeth (Costa collection) can be referred to small theropod dinosaurs, possibly dromaeosaurids (see BUFFETAUTet ai., 1986, for an earlier report from Villeveyrac). Iguanodontia incertae sedis cf. Rhabdodon priscus MATHERON,1869 Cranial and post-cranial ornithopod remains from Villeveyrac are provisionally referred to as cf. Rhabdodon priscus. This material includes four teeth, dorsal and caudal vertebrae, a humerus and a fragmentary femur.

Fig. 6. cf. Rhabdodon priscus (Early Campanian, Villeveyrac, cast MDE-Vil-14); dorsal vertebra in cranial (A), lateral (B) and caudal (C) views; scale bar: 5 em. Drawings by GUY LE Roux (Musee des Dinosaures, Esperaza). A maxillary tooth (COSTACoil., noad-10) has a rostro-caudallength of 20 mm; it bears 19 vertical ridges. Three other teeth (VALENTIN collection; uncatalogued) are worn dentary teeth. A large dorsal vertebra (VALENTINcollection; uncatalogued; cast MDE-Vil-14) is remarkable for its very tall neural spine (Fig. 6). Three amphicoe1ous caudal centra are preserved (COSTAcollection). Ventrally, they bear a single chevron facet anteriorly and posteriorly, as in Rhabdodon. Telmatosaurus and other hadrosaurs bear two well-separated facets anteriorly and posteriorly. A proximally incomplete humerus (VALENTINcollection, uncatalogued; cast MDE-Vil-13) shows a laterally projected delto-pectoral crest. The above-mentioned teeth can be referred to the genus Rhabdodon MATHERON,1869, because of their peculiar morphology, and the postcranial ornithopod remains from the Villeveyrac locality in alllike1ihood also belong to that taxon. This ornithopod is known from the Campanian of Austria and the Maastrichtian of Spain, France and Romania (see BRINKMANN1988). The very tall neural spine of the dorsal vertebra is reminiscent of undescribed Rhabdodon priscus vertebrae from the Maastrichtian of Provence (MECHIN collection) and in this respect the Villeveyrac specimen differs from the Transylvanian Rhabdodon

vertebrae which have low neural spines (NOPCSA). Together with other osteological differences (del to-pectoral crest of the humerus cranially oriented in the Romanian species versus laterally projected in the specimens from Southern France), this suggests that the material from Romania cannot be referred to the same species (LE LOEUFF 1992). We refer the material from Romania to Rhabdodon robustus (NOPCSA 1900) on the basis of the above-mentioned differences. The bones from Provence can be referred to Rhabdodon priscus MATHERON 1869. The very high neural spine of the Villeveyrac vertebra and the laterally projected del topectoral crest of the humerus suggest that it is related to the Maastrichtian species from Provence. Pending the completion of a study of new Rhabdodon material from southern France, we refer to the Villeveyrac material as cf. Rhabdodon priscus. Ankylosauria Nodosauridae indet. Two osteoderms (COSTA collection, uncatalogued and MDE-ViI-01) and three teeth (VALENTIN collection, uncatalogued; cast MDE-Vil-17) belong to an ankylosaur. The well-developped cingulum of the teeth IS reminiscent of the family Nodosauridae (ASTIBIA et al. 1990). Conclusions The Villeveyrac fauna is the most complete Early Campanian terrestrial fauna hitherto recovered from western Europe. Although its composition is not unexpected in view of what is already known from other European sites, it has various palaeoecological and biostratigraphical implications. The occurrence in the same beds of remains of palaeobatrachids, lepisosteid and sparids is noteworthy. This association is very interesting from an ecological point of view since the Palaeobatrachidae and Lepisosteidae are strictly indicative of a freshwater environment while the Sparidae suggest shallow marine or brackish deposits. This probably implies an estuarine environment or the existence of pools or lagoons near the shoreline resulting in salinity changes, which in turn caused changes in the vertebrate fauna. These changes may be explained by various phenomena, such as connections with both the sea and a fluvial system, cyclical drought and rainfall causing salinity changes, or cyclical tide-related changes. Other alternative explanations for the association of Lepisosteidae and Sparidae may be found, but in any case a marginal environment close to the shoreline seems most likely. On the basis of the invertebrate fauna, FREYTET (1970, 1971) came to similar conclusions

about the coexistence and occasional mingling of freshwater and brackish assemblages in the basal series of the Villeveyrac basin. Among the main peculiarities of the Villeveyrac fauna is the absence (so far) of titanosaurid dinosaurs, which are the dominant herbivorous dinosaurs in the Early Maastrichtian of western Europe. This absence is also remarkable in the other Early Campanian localities in Europe. Two hypotheses may explain this fact: either a Late Campanian immigration of titanosaurids from Africa (Late Campanian titanosaurids are known from marine deposits in southwestern France: see PLATEL1989) following an earlier, possibly Cenomanian extinction of this group in Europe (see LE LOEUFF1993, for a review of European titanosaurids) or an ecological explanation suggesting that titanosaurids were not present in lagunar environments such as that assumed for the Villeveyrac deposits. Recent unpublished footprint discoveries in Catalonia (A. MARTINEZ, pers. commun.) show that titanosaurids were sometimes common in lagunar environments in the Late Maastrichtian, which seems to be in favour of the first hypothesis of a "titanosaurid hiatus" (see LUCAS& HUNT 1989) during the early part of the Late Cretaceous in Europe. Admittedly, however, the available data are too scanty to warrant a definitive answer to that question.

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