LATE CRETACEOUS VERTEBRATES FROM THE …

1 ACTA PALAEONTOLOGICA ROMANIAE V. 5 (2005), P. 39-48. late CRETACEOUS VERTEBRATES FROM THE SAINT-CHINIAN AREA. (SOUTHERN FRANCE): A REVIEW OF PREVIOUS RESEARCH AND AN UPDATE ON. RECENT FINDS. Eric BUFFETAUT1. Abstract: This paper traces the history of research on the late CRETACEOUS VERTEBRATES of the Saint- Chinian area of southern France since the first report of dinosaur bones by Gervais in 1877. A number of researchers, including Miquel, Dep ret, Nopcsa and Lapparent, were involved in the early studies on this faunal assemblage, from the 1890s to the 1950s, but the known number of taxa remained low because specimens were recovered almost exclusively by surface collecting. A faunal list showing a significantly higher diversity is presented, on the basis of recent work involving systematic excavations and screenwashing. This list includes actinopterygians, coelacanths, amphibians, turtles, squamates, crocodilians, pterosaurs, dinosaurs, birds and mammals.

2 Fossil eggs are also common. On the basis of its composition, the Saint-Chinian vertebrate fauna is referred to the Early Maastrichtian. Keywords: late CRETACEOUS , Vertebrata, Southern France, Fauna. INTRODUCTION assignment being difficult because of the lack of stratigraphically useful fossils, marine ingressions The region around the small town of Saint- or radiometric dates. Until the 1960s, most authors Chinian, in the western part of d partement considered them as belonging to the latest stage of H rault, in southern France ( ), has been the CRETACEOUS , for which various names were known to yield dinosaur remains since 1877. In the used (Danian, Maastrichtian, or Rognacian the late 1890s, more systematic collecting resulted in latter being a local stage name defined in the identification of several dinosaur taxa, including Provence). Comparisons with the similar the first titanosaurids to be reported from the late stratigraphic succession in Provence have often CRETACEOUS of Europe.

3 Despite further work in the been attempted (hence the use of the name middle decades of the 20th century, the late Rognacian), but the exact ages of the non-marine CRETACEOUS vertebrate assemblages of the Saint- late CRETACEOUS formations of Provence are still Chinian area remained poorly known until somewhat uncertain. As noted by Miquel (1897), systematic excavations at several sites were the limestones at the top of the CRETACEOUS series begun in the 1990s. The present paper includes a in the Saint-Chinian area contain freshwater review of the work done since the 1870s, followed molluscs also known in the Rognac Limestone of by an update on the results acquired during the Provence, and in the light of current knowledge, last decade (for a more extended, semi-popular this certainly suggests a Maastrichtian, possibly account, see Buffetaut, 2005). late Maastrichtian, age for them. However, most of the fossil VERTEBRATES from that area come from the GEOGRAPHICAL AND GEOLOGICAL SETTING underlying sandstones and clays, which can be early Maastrichtian or older.

4 Freytet (1965). The Saint-Chinian area is located NW of the reported the occurrence of brackish molluscs city of B ziers, between the coastal plain which suggesting a relatively early age (possibly as early borders the Mediterranean and the Palaeozoic as Turonian, more probably Campanian), on the massif of the Montagne Noire. Its hilly relief is the basis of correlation with similar assemblages from expression of a complex geological structure, the Provence. However, this brackish assemblage was area having been heavily folded and faulted during found at a single locality (Fontanche, near the the Cenozoic, which resulted in what is known to village of Cruzy) in beds which in fact underlie the French geologists as the Cha non de Saint- vertebrate-bearing sandstones and clays, so that Chinian . its age, which itself is uncertain, says little about In that area, the non-marine late CRETACEOUS is that of the overlying vertebrate-bearing red beds.

5 Well represented by red beds consisting of By comparison with other assemblages from conglomerates, sandstones, and clays and by southern France, the composition of the vertebrate freshwater limestones, sometimes known as the fauna from the Saint-Chinian area seems to Gr s reptiles , overlain by basal Tertiary red indicate a late Campanian to early Maastrichtian clays. Vertebrate remains occur, sometimes in age. The eggshell assemblage from that area is abundance, in all these facies, the most productive reported to suggest an early Maastrichtian age localities being in the clayey layers. The exact age (Garcia & Valentin, 2001-2002). of the late CRETACEOUS vertebrate-bearing beds in the Saint-Chinian area has been a matter of discussion (see below for a review), a precise age 1. CNRS (UMR 5125, Pal oenvironnements et Pal obiosph re), 16 cour du Li gat, 75013 Paris, France. Eric BUFFETAUT. Fig. 1. Map of part of southern France showing the Saint- Chinian area (darker shading) described in the present paper.

6 A BRIEF HISTORY OF RESEARCH ON late He was thus the first to mention that fossils similar CRETACEOUS VERTEBRATES IN THE SAINT- to the large reptiles from Provence had also been CHINIAN AREA found at Saint-Chinian (and also at Esp raza, in the Aude valley an area where highly productive During the 19th century, vertebrate remains, dinosaur localities were discovered in the 1980s;. including both bones and eggs, were discovered in see Buffetaut et al., 1989). Unfortunately, no non-marine sediments of late CRETACEOUS age in detailed descriptions were provided by Gervais, various parts of southern France, from Provence to who died soon thereafter, and these specimens the foothills of the Pyrenees (Buffetaut et al., seem to have been lost. 1993). However, few detailed descriptions of that After Gervais's brief mention, little seems to material were published at the time, Philippe have been done about the fossil VERTEBRATES from Matheron's 1869 paper on fossil reptiles from the Saint-Chinian area until the 1890s, when an Provence being an exception (Matheron, 1869).

7 Enthusiastic and gifted local researcher, Jean While Jean-Jacques Pouech's similar discoveries Miquel (1859-1940), became interested in them. in Ari ge (Pouech, 1859) had been ignored by the Miquel, who had studied law at the university of French scientific community (Buffetaut, 1992; Le Toulouse, was a landowner in the wine-making Loeuff, 1992; Buffetaut & Le Loeuff, 1994), area around Saint-Chinian. He developed a keen Matheron's discovery of what turned out to be interest in the geology, palaeontology and large eggs (made before Pouech's, but announced archaeology of that region, and thoroughly later) at Rognac, near Marseille, attracted the explored the region around his estate at Barroubio, attention of Paul Gervais (1816-1879), then at the publishing a number of papers in local scientific Comparative Anatomy Laboratory of the Paris journals. His main geological and palaeontological Museum. In 1877, Gervais published two identical contributions were on the Palaeozoic strata of the papers on the microstructure of eggshells, both Montagne Noire (Alvaro & Vizca no, 2002), to the recent and fossil (Gervais, 1877a,b).

8 This was a north of the CRETACEOUS outcrops, but he also landmark in the study of fossil eggs, showing how published on the Mesozoic stratigraphy of the different microstructural types could be region (Miquel, 1897, 1905) and mentioned the distinguished and tentatively used for the occurrence of bones of large reptiles at several systematic identification of fossil eggs (Buffetaut & levels in what he called the fluvio-lacustrine Le Loeuff, 1994; Taquet, 2001). In that paper, CRETACEOUS series . He correctly referred these Gervais also reviewed what was then known about strata to the Rognacian , a local stage defined in the late CRETACEOUS dinosaurs of southern France. Provence and now considered as an equivalent of 40. late CRETACEOUS VERTEBRATES from the Saint-Chinian area (southern France). the Maastrichtian. Although Miquel did not attempt observations, he thought that the dinosaurs of the to precisely identify the late CRETACEOUS vertebrate Saint-Chinian area occurred at two distinct levels, remains he discovered, he provided interesting some being from sandstones at the base of the details about their occurrence.

9 In 1897, he Rognacian limestones, whereas most of them tantalisingly noted that twenty years before, a (notably at Castigno) came from clays which he reptile skeleton had been found by a Mr Salles thought overlay the limestones, and were an when digging a canal near Saint-Chinian and had equivalent of the Vitrolles Clays of Provence. been studied by Paul Gervais , but had Dep ret considered that the presence of dinosaurs subsequently been lost (as mentioned above, in these strata demonstrated their terminal Gervais never described reptile material from CRETACEOUS age, and he referred them to the Saint-Chinian in any detail). Miquel himself found uppermost part of the Danian (then considered as reptile bones in abundance in the late CRETACEOUS the last stage of the CRETACEOUS ). Another sandstones and clays. They were especially interpretation was possible, though, as expressed abundant in the Castigno gully, near the village of by Emile Haug in his famous Trait de G ologie Villespassans, from which he often brought back (Haug, 1908-1911): the presence of dinosaur loads of 10 to 15 kg of fossil bones (Miquel, 1897).

10 Remains in an equivalent of the Vitrolles Clays, He purposely left some large bones as witnesses widely considered as basal Tertiary (Montian) in in the sandstones forming the banks of the gully; age, could also be taken to indicate that dinosaurs some of them were rediscovered by our group in had survived into the Tertiary. Haug favoured this the 1990s. interpretation, which he thought was supported by Jean Miquel lacked the expertise and the the purported occurrence of dinosaurs in the comparative material needed to identify the late Tertiary of Patagonia, as reported by Ameghino. CRETACEOUS vertebrate remains he found in the Ameghino's claims were later shown to be founded Saint-Chinian area and wisely refrained to do so. on faulty stratigraphical interpretations and The first precise identifications were published in misidentifications of ziphodont crocodilian teeth. As 1900 by Charles Dep ret (1854-1929), a professor to the supposedly Tertiary dinosaurs from Saint- of geology at the University of Lyon and expert on Chinian, the problem was solved by Lapparent, fossil VERTEBRATES .