french dinosaurs: the best record in

ll4odern Geology, 1991, Vol. 16, pp. l7 12
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FRENCH DINOSAURS: THE BEST RECORD IN
EUROPE?
ERIC BUFFETAUT. GILLES CUNY and JEAN LE LOEUFF
CNRS, Laboratoire de Paléontologie des Vertébrés, Université Paris VI,
4 place Jussieu, 75252 Paris Cedex 05, France
(Received December 29, 1990)
Although dinosaur remains from France were described
as early as the beginning of the 19th century, the
importance of the French dinosaur record has been generally underrated, partly because of a relative lack
of interest on the part of French palaeontologists. A review of the available data (including historical
aspects) shows that, although few spectacular finds have been reported, the French record is stratigraphically fairly complete, with finds ranging in age from Late Triassic to terminal Cretaceous. Fairly abundant
remains have been found in the Upper Triassic, Upper Jurassic and Upper Cretaceous, but deposits from
other periods have also yielded important specimens. In terms of stratigraphic completeness, the French
dinosaur record seems to be the best in Europe and renewed studies can contribute importantly to an
understanding of dinosaur faunal succession on that contlnent.
KEY WORDS: Dinosauria, France, fossil record, stratigraphy, historical review.
INTRODUCTION
France does not readily come to mind when one thinks of places in the world where
dinosaurs have been found in abundance. Even within the narrower framework of
European countries, the historical discoveries made in England during the early
nineteenth century and the spectacular finds of lguanodon at Bernissart in Belgium
or of Plateosaurus aI Trossingen and Halberstadt in Germany certainly overshadow
the French record of mostly isolated finds. Nevertheless, a review of French
dinosaurs reveals that they are much more numerous than is usually realised (Figure
1), and that the French record spans almost the whole of dinosaur history, from the
Late Triassic to the end of the Cretaceous. Our aim in this paper is to review briefly,
but as thoroughly as possible, the record of dinosaur finds from France. Considering
the systematic uncertainties about many of the specimens, a stratigraphic order has
been chosen. Whenever it was suitable, historical details have been provided. The
next section is a brief outline of dinosaur research in France since the beginnings of
scientific palaeontology.
THE DISCOVERY OF FRENCH DINOSAURS:
A HISTORICAL SKETCH
As pointed out by Taquet (1984), there may have been dinosaur remains among the
fossil bones found by Abbé Dicquemare in the Jurassic beds in the vicinity of Le
Havre, although his descriptions publishedinlTT6 are not sufficient to confirm this.
The earliest uncontrovertible description and illustration of dinosaur bones from
France was published by Cuvier in 1808, in a paper devoted to fossil bones from the
Jurassic of Honfleur, in Normandy. Among this material, which he referred to
l1
E. BUFFETAUT, G. CUNY AND J. LE LOEUFF
18
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I Geographical distribution of main French dinosaur localities (including both footprint sites and
skeletal finds). Triangles: Triassic. Squares: Jurassic. Circles: Cretaceous. B: Brest, Bo: Bordeaux, L:
Lyon, M: Marseille, P: Paris, S: Strasbourg, T: Toulouse.
Figure
crocodiles, were theropod vertebrae (Figure 2). After a visit to Buckland in Oxford
in 1818, during which he was shown the bones which were to be described as
Megalosaurus, Cuvier did realise that some of his vertebrae from Honfleur were
similar to those of the large reptile from Stonesfreld, as shown by his remarks on this
point in the second edition oThis Recherches sur les ossemens fosslles (Taquet, 1984).
However, Cuvier did not elaborate much on the French material in his possession,
although he was later involved in the identification of dinosaur material from
England, including remains of lguanodon submitted to him by Mantell (Taquet,
1984; Buffetaut, 1987).
An important early find of dinosaur remains in France was that of a partial
in the Bathonian "Pierre de Caen" in the 1830s. J. A.
skeleton of a Theropod
Eudes-Deslongchamps, who described the specimen as Poekilopleuron Bucklqndii
in 1838, has recounted in some detail the complicated and sometimes rather farcical
circumstances of the discovery (Eudes-Deslongchamps, 1B3B; see also Buffetaut,
1983. 1987).
t9
FRENCH DINOSAURS
.t
#1,{ y'r',x"/""u,,:,...}1,.,'.,cii . dl.
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Figure
2
The oldest published illustration of French dinosaur remains: theropod vertebrae from the
Jurassic of Normandv described by Cuvier (1808) as those of peculiar crocodiles. From Cuvier (1808).
The first palaeontologist to have applied Owen's name, Dinosauria, to French
specimens seems to have been Paul Gervais, who in 1848-52 figured several
specimens under that general name, in a book in which he also erected the families
Megalosauridae and Iguanodontidae. The material then available to Gervais was
scanty. During the second half of the nineteenth century, however, dinosaur remains
were found in many parts of France, in rocks ranging in age from Late Triassic (with,
notably, the Plateosaurus remains from the Poligny region described by Pidancet and
Chopard, 1862, as Dimodosauras) to Late Jurassic (with, for instance, the remains
from Boulonnais described by Sauvage in a series of papers) and Late Cretaceous
(Matheron's paper of 1869 marking the beginning of the study of the rich localities of
Provence). F{owever, these various finds did not always attract the attention they
deserved beyond the regional level. This may have been due to a relative lack of
interest on the part of Parisian palaeontologists. Albert Gaudry, who for many years
was the head of the palaeontology department of the Muséum national d'Histoire
naturelle, was an expert on fossil mammals. Although he did devote more than 20
pages of his Enchqînements du monde animal (1890) to dinosaurs, he does not seem
to have been very keen on excavating dinosaur localities himself. When Abbé J. J.
Pouech wrote to Gaudry in 1880 to inform him of the discovery of a remarkably rich
vertebrate locality in the Upper Cretaceous of the Pyrenees, there was apparently no
reaction from Gaudry, who only read Pouech's letter at a meeting of the Société
géologique de France. This did not prevent Gaudry from (unsuccessfully) trying to
acquire dinosaurs discovered by others, including one of the lguanodon skeletons
from Bernissart (see Buffetaut, 1987) and the stegosaur found in the Kimmeridgian
of Normandy by Savalle and Lennier. Many good opportunities for interesting finds
and productive excavations were certainly lost because of this relative lack of
interest, at a time when many quarries and outcrops which today no longer exist were
still accessible.
Gaudry's successor at the Paris Museum, Marcellin Boule, is mainly remembered
for his work on fossil man, and his able assistant Armand Thevenin, who
was
20
E. BUFFETAUT, G. CUNY AND J. LE LOEUFF
interested in reptiles, including dinosaurs, died in 1918 as a result of experiments on
poison gas. Despite some important chance discoveries, such as that ol the Oxfordian dinosaurs of Damparis (Jura) in 1934, very little was published on French
dinosaurs between 1914 and 1945. In 1939, however, A. F. de Lapparent conducted
systematic excavations in the upper Cretaceous of Fox-Amphouf (var), and in 1947
he was able to publish an important monograph on the Late Cretaceoué dinosaurs of
southern France. Until his death in 1975,Lapparent was the main expert on French
dinosaurs, on which he published a short review paper in 1967. Although some
important material (including the forgotten lguanodon material of the Saiù-Dizier
region in the eastern Paris Basin) was rediscovered in museums by Lapparent during
the 1950s and 1960s, few significant new finds of skeletal remaini of àinosaurs weré
made (exceptions being the "Rhaetian" theropod found at Airel (Manche) by
Larsonneur and the callovian stegosaur found at Argences (calvados) by R. Brunj,
but important work was done on the Late Cretaceous eggshells of southern Franée
(notably by Dughi and Sirugue) and on several footprint iites. Renewal of interest in
French dinosaurs in
J!9 1970s was spurred by the discovery of the canjuers
compsognathus. rn 1979, excavations were started again at Fox-Amphoux. Êven
more recently, following discoveries by amateur palaeontologists, large-scale excavations have been started by our group both in the Upper Cretaceous of the Upper
Aude Valley and in the Upper Triassic of Jura.
A SURVEY OF THE RECORD
As explained above, a stratigraphic arrangement seems to be the best way to present
the record of French dinosaurs. This should not be considered as a comprehensive
review, because some undescribed material could not be included. Althoush footprint and eggshell sites have been mentioned, the emphasis has deliberately been put
on the finds of skeletal remains.
l.
Middle Triassic
No skeletal remains of dinosaurs have hitherto been found in the Middle Triassic of
France. The vertebrae from the Muschelkalk of Lunéville (Meurthe-et-Moselle)
referred ro Thecodontosaurus latespinatus Huene, 1908 by Huene (1908) and Corroy
(1928) have been shown to belong to Tanystropheus by Wild (1973)
Footprints from several Middle Triassic localities on the southern and eastern
edges of the French Central Massif (distribution map in Demathieu, 1989) have been
referred by Courel et al. (1968) to the ichnogenera Anchisauripus, Coelurosaurichnus and Grallator. According to Demathieu (1989), these tridactyl "dinosauroid"
tracks were made by early dinosaurs, as early as the Middle Anisian. A confirmation
by the discovery of skeletal remains of true dinosaurs in contemporaneous sediments
in Europe would be welcome.
2. Late Triassic
The earliest uncontrovertible skeletal remains of dinosaurs hitherto reported from
France are from the Late Triassic continental deposits of the eastern part of the
country, from Lorraine in the north to the southern part of the Jura mountains in the
south.
FRENCH
DINOSAURS
21
one of the first important discoveries was reported in 1862 by pidancet and
chopard (see also chopard, 1883), who described remains of a "gigantic saurian"
found in the "marnes irisées" of the vicinity of Poligny (Jura). They proposed the
name Dimodosaurus poligniensis for this animal (Figure 3) , which was said to be very
abundant in the local Keuper, remains of five individuals having been found in thê
railway cutting at Chassagne near Poligny, and those of two individuals in the Villette
cutting near Arbois (Jura). Other Dimodosaurus localities mentioned by Pidancet
and Chopard are Domblans (Jura) and Beurre, near Besançon (Doubs). The Poligny
specimens of Dimodosaurus were split between the genera Plateosaurus and Gresslyosaurw (a junior synonym of Plateosqurus according to Galton, 1985) by Huene
in 1908. The material from the Poligny region, which belongs to the collections of the
Dl
lltlnos.tu ttus
pilt ci'l:x.sts
(Aei,p<.rù'1s Eoupos )
llal,rncnt du. ;yuLao. 1 ,iz. ai /a yra,rLar rv/c)
Figure 3 An early reconstruction of part of the skeleton of a Triassic dinosaur from eastern France:
" Dimodosaurus poligniensis" (now referred to Plateosaurus) from the Upper Triassic of Poligny (Jura), as
illustrated by Chopard (1883).
22
E. BUFFETAUT, G. CUNYAND J. LE LOEUFF
Poligny museum, is currently being revised by one of us (G.C.); it comprises well
preserved postcranial elements of several skeletons. Since the discoveries of Pidancet and Chopard, the region of the Jura mountains has yielded a number of
additional plateosaur remains. As early as 1867, Ogérien reported a find from
Feschaux (Jura), which was identified by Huene (1908) as Gresslyosaurus. A well
preserved Plateosquru.ç foot found at Violot (Haute-Marne) at the end of the
nineteenth century belongs to the Langres Museum; although it has been mentioned
under various names (ThecodontosaLtrus, Zanclodon) by several authors (Thiéry,
1910; Corroy,1934), it has never been described and is currently being studied by
one of us (G.C.). A few Plateosqurus bones from Boisset (Jura) were described in
1967 by Theobald et al. Finally, excavations have been started in 1990 by G. Cuny
and J. M. Mazin at a rich new Plateosaurus locality at Lons-le-Saunier (Jura). All the
above-mentioned Plateosaurus material comes from beds referrable to the Keuper,
and is probably Norian in age, although uncertainties as to the exact stratigraphic
origin of many of the older finds remain.
Some supposedly "Rhaetian" localities have also yielded dinosaur remains. If, as
accepted by many authors, the "Rhaetian" is only an equivalent of the upper Norian
(see discussion in Buffetaut and Wouters, 1986), these specimens should be considered as Norian in age. Teeth from Moissey (Jura), described as Megalosaurus
obtusus by Henry (1876), were referred to Plateosaurus by Huene (1908); Galton
(1985) has rightly considered them as those of an indeterminate archosaur. Similarly,
teeth from Provenchères-sur-Meuse (Haute-Marne) describe d as Thecodontosaurus
elizae by Sauvage (1907) were also referred to Plateosaurus by Huene (1908).
fïom Provenchères in the collections of the University
of Paris, under study by G. Cuny, confirm the occurrence of prosauropod
Undescribed material
remains.
Teeth found by Dumortier in the "marnes irisées" at Le Chappou, near SaintRambert (Ain), were described by Gervais (186i, 1869) and referred by him to
Thecodontosaurus. They may be considered as those of an indeterminate prosauroood.
Teéth of Plateosaurus and of a small theropod (possibly a procompsognathid),
now kept at the Institut royal des Sciences naturelles de Belgique (Brussels), have
been described by Buffetaut and Wouters (1986) from Saint-Nicolas-de-Port
(Meurthe-et-Moselle). The locality is famous for having yielded mammal teeth and
has often been referred to the Rhaetian. However, the composition of its amphibian
and reptile fauna has led Buffetaut and Wouters (1986) to place it in the Middle
Norian, at roughly the same level as the German Knollenmergel. Current work by G.
Cuny on an unpublished coliection from Saint-Nicolas-de-Port seems to confirm
this age assignment.
A
vertebra from the "Rhaetian" of Pierreclos (Saône-et-Loire) referred to
Plateosaurus by Lapparenl (1961) and kept in the Paris Museum in fact belongs to a
sauropterygian (G. Cuny, personal observation).
A few footprint sites with "dinosauroid" tracks have been recorded in the Upper
Triassic of southern France. The Late Triassic (Keuper) site at Anduze (Gard)
contains tridactyl footprints associated with enigmatic "elephant-like" ones (Ellenberger, 1965). According to l-apparent and Montenat (1867), the lower footprintbearing beds of the famous site of Le Veillon , on the Atlantic coast of Vendée, are
likely to be "Rhaetian" in age, whereas the upper ones are referred to the Hettangian. More than 1000 footprints have been found at Le Veillon: they include the
tridactyl ichnogenera Grallator, Eubrontes, Saltopoides, Anatopus
Talmonlopus.
and
FRENCH DINOSAURS
3. Early Jurassic
Few dinosaur remains have been reported from the Lower Jurassic of France. In
1855, serrated reptile teeth from the Hettangian of Hettange (Moselle) were referred
by Terquem to Megalosaurus. rn 1859, some of these teeth were figured and
described by Gervais as Megalosaurus terquemi. rnlg6l , Lapparent referred to this
material as Gressylosaurus (sic) terquemi.In 1954, however, Huene and Maubeuge
described similar teeth from the Hettangian of Hettange and referred them tJa
parasuchian (phytosaur). This identification seems correct (it is the only record of a
post-Triassic phytosaur), and it is likely, from the figures and descriptions, that the
teeth described by Terquem and Gervais also were phytosaur teeth.
An important apparently Early Jurassic fossil has been found in Normandy, in a
quarry near the village of Airel (Manche). The specimen, an incomplete skeleton
kept at the university of caen, comprises many vertebrae and a large part of the
pelvis, as well as a tooth; it was described in 1966 by Larsonneur and Lapparent, who
identified it as Halticosaurus sp. The bones have now been prepared more fully and a
revision by G. Cuny and P. M. Galton is under way; preliminary comparisons suggest
that the Airel dinosaur may be allied to Liliensternus Welles, 1984. The non-marine
Airel beds, which have yielded the specimen, were first thought to be "Rhaetian" in
age, but a palynological study by Médus (1983) suggests that they may in fact be
Hettangian or early Sinemurian in age. Palynological studies on the matrix surrounding the bones have been conducted (by R. Rauscher and J. Doubinger, Strasbourg)
and suggest an Early Jurassic, rather than Triassic, age.
A tooth referred 1o Megalosaurus cloacinus Quenstedt, from the Lower Hettangian of the calcaire de valognes at valognes (Manche), has been mentioned by
Rioult (1978a) as having been destroyed by an air raid on the University of Caen in
t944.
Early Liassic dinosaur trackways u,eré' reported by Thaler (1962) from the
Hettangian of Lodève (Hérault) and Saint-Laurent-de-Trèves (Lozère). These
tridactyl footprints were attributed to fairly large theropods. Small tridactyl footprints were reported from the "Lotharingian" of Séverac-le-Château (Aveyron) by
Ellenberger and Fuchs (1965).
4. Middle Jurassic
The Bathonian and Callovian rocks of Normandy have yielded a fairly large number
of dinosaur specimens. They occur in marine deposits and can be interpreted as
of floated carcasses. There are also a few reports of Middle Jurassic
dinosaurs from other parts of France.
remains
Bathonian Isolated theropod teeth referre dTo Megalosaurus are reported from the
Early Bathonian Calcaire d'Ecouché of Ecouché, Orne (Rioult, 1978a). One tooth
from the Lower Bathonian of Les Aucrais (Calvados) was attributedto Megalosaurus bucklandi by Mercier (1937), who also mentioned vertebrae from the same
quarry.
As early as 1828, A. de Caumont reported the occurrence of Megalosaurus
remains in the Calcaire de Caen, or "Grande Oolithe," now known to be Middle
Bathonian in age. One of the most complete theropod skeletons ever found in France
was discovered in the Calcaire de Caen at Caen (Calvados) in the 1830s. and
described by J. A. Eudes-Deslongchamps (1838) as Poekilopleuron bucklandi. lt
was later renamed Megalosaurus poekilopleuron by Huene (\932). The specimen,
24
E. BUFFETAUT, G, CUNY AND J. LE LOEUFF
which comprised vertebrae, ribs, gastralia and many elements of the limbs, was
destroyed by an Allied air raid on the University of Caen in 1944. An isolated
theropod tooth from the "Grande Oolithe" of Quilly (Calvados) mentioned by
Gervais (1859) suffered the same fate.
Additional theropod remains from the Calcaire de Caen are mentioned by Rioult
(1978a) as Megalosaurus sp.
In west-central France, the Bathonian of Saint-Gaultier (Indre) has yielded two
isolated dinosaur teeth described by Sauvage (1900); one was a theropod tooth which
he referred to Megalosaurus bucklandi,lhe other was a spatulate sauropod tooth
which was attributed to Pelorosaurus.
Callovian The Callovian of Normandy has yielded a variety of dinosaur bones. The
exact geographical and stratigraphic origin of some of the specimens in the older
collections may be uncertain because of possible confusions with Kimmeridgian
specimens lrom the same general area.
One of the most complete dinosaur specimens hitherto found in the Callovian of
France is the partial stegosaur skeleton found by R. Brun in a clay-pit at Argences
(Calvados) in 1955 (Hoffstetter,I95T; Hoffstetter and Brun, 1956, 1958; Galton,
Brun and Rioult, 1980). The specimen, from the Lower Callovian, is now kept in the
Natural History Museum in Le Havre. It was referred to the new genus Lexovisaurus
by Hoffsteter in 1951, and Galton et al. (1980) provisionally referred it to L.
durobrivensis (Hulke).
It
comprises 25 vertebrae, 12 hemal arches,
rib
fra-
gments, the left humerus, the right femur, tibia, fibula, astragalus and calcaneum,
plus a large right parasacral spine. A redescription is being prepared by
P. M. Galton.
A number of theropod remains have also been obtained from the Callovian of
Calvados (lists have bèen provided by Huene ,1926 and Rioult, 1978a). Most of the
material is Late Callovian in age. However, Lydekker (1888) mentions "the proximal extremity of a very large left tibia" referred to Megalosaurus bucklandi, in the
British Museum, "probably from the Fuller's Earth near Caen." This would make it
Bathonian, but Huene (1926) refers it to the Lower Callovian. He also places in the
Lower Callovian the distal part of a very large femur and a phalanx, kept at the
University of Tùbingen; the specimens are said to come from the Vaches Noires near
Dives. However, the only Callovian levels (the "Marnes de Dives," well-known for
their fossil vertebrates) at the base of the Vaches Noires cliffs, between Villers-surMer and Houlgate, on the Calvados coast, are Late Callovian in age (see Rioult,
1978b), and Huene's age assignment seems doubtful.
Huene (1926) lists a variety of "megalosaurian" bones from the Upper Callovian
of the Vaches Noires and neighbouring areas (Dives, Beuzeval, etc.), consisting
mainly of vertebrae and limb bones kept in Paris (Natural History Museum and
Ecole des Mines), Caen (University Museum) and London (British Museum). The
material kept in Caen was destroyed in 1944.
A specimen of special importance is the theropod braincase described by Piveteau
(1923) as belonging to Streptospondylus cuvieri. The fossil comes from the Vaches
Noires, near Dives, and is of Late Callovian age (Rioult, 1978a). It was called
Eustreptospondylus divesensis by Walker (1964), and then restudied by Taquet and
Welles (1917) , who used it as the holotype of the new taxon Piveteausaurus divesensis
(however, the possibility remains that it could actually belong to Megalosaurus,in
which the braincase is not known with certainty; the braincase from the Bathonian of
England described as Megalosaurus by Huene in 1906 may well be that of a
sauropod). As rightly pointed out by Taquet and Welles, some postcranical elements
FRENCH
DINOSAURS
25
described by Piveteau (1923) together with the braincase from Dives in fact probably
come from younger deposits near Honfleur.
A maxilla fragment from the Upper Callovian of the Vaches Noires near Villers,
found in 1987, is currently being described as Megalosaurus sp. by E. Buffetaut, G.
Pennetier and E. Pennetier.
Postcranial remains of a theropod have recently been reported from the Callovian
of Puits-de-1a-Brême (Doubs) by Broin et al. (1997).
5. Late Jurassic
Late Jurassic dinosaurs are known from various parts of France. Some occur
as
floated carcasses in marine deposits, whereas others have been found in non-marine
sediments (this reflecting a trend toward more continental deposition through the
Late Jurassic).
Oxfordian A few dinosaur remains have been reported from the marine Oxfordian
levels of the Vaches Noires cliffs (Calvados). They include an undescribed large
theropod vertebra kept in the palaeontological museum at Villers-sur-Mer
(mentioned by Rioult, 1978a).
Isolated dinosaur teeth from the Late Oxfordian fluvio-marine Sables de Glos, at
Cordebugle near Lisieux (Calvados), have been described by Buffetaut et al. (1985).
They include theropod teeth and one spatulate sauropod tooth resembling those of
the Camarasauridae.
A tooth from the Sables de Cherré (Sarthe) was referred by Bigot (1904) to
Pleurocoelus, a sauropod genus originally described by Marsh from the Lower
Cretaceous of Maryland. The specimen never was described or figured, and seems to
be lost. The Sables de Cherré, of Late Oxfordian age, are a southern equivalent of
the Sables de Glos, and the tooth mentioned by Bigot mav have been similar to the
above-mentioned one from Cordebugle.
A remarkable dinosaur discovery was made in 1934 in a limestone quarry at
Damparis, Jura (Dorlodot, 1934), when a large part of a sauropod skeleton
associated with seven theropod teeth was found in a marly lens intercalated in marine
limestone. Although Lapparent, who described the dinosaur remains in 1943,
referred them to the Kimmeridgian in 196l , recent stratigraphic studies show that
they belong to the Late Oxfordian (Buffetaut, 1988). Lapparent identified the
sauropod remains as Bothriospondylus madagascariensis, which seems highly questionable ; a revision of the material (in the Paris Museum) is desirable. FIe referred
the theropod teeth to Megalosaurus insignis (a species based on an isolated tooth,
and hence of doubtful validity: see below). The Damparis dinosaur remains have
long been considered as those of carcasses buried in marine sediments after floating
from a distant land area. However, a careful analysis of the circumstances of the find
and of the association of a sauropod skeleton with theropod teeth leads to the
conclusion that the sauropod carcass was preyed upon by theropods ln sl/u during an
episode of emersion, also suggested by other data (Buffetaut, 1988).
"Megalosaurus nicaeensis," described by Ambayrac (1913) on the basis of a large
jaw from the Oxfordian of La Turbie (Alpes-Maritimes), is in fact a large pliosaur
(Buffetaut, 1982).
A few remains from Late Oxfordian ("Rauracian") lithographic limestones at
Hadainville, near Verdun (Meuse), have been referred to dinosaurs by Huene and
Maubeuge (1954). They include a laterally compressed tooth with a forked posterior
cutting edge, interpreted as a possible "coelurosaurian," a proximal rib fragment
referred to a sauropod, and a fragmentary pubis referred to a "camptosaurid."
26
E. BUFFETAUT, G. CUNY AND J. LE LOEUFF
Lapparent (1967) mentions Megalosaurus sp. in the Oxfordian
(Vienne). We have not been able to confirm this report.
of
Poitiers
Kimmeridgian Dinosaur remains have been found in relative abundance in the
marine Kimmeridgian of Normandy, near Le Havre (Seine-Maritime) and Honfleur
(Calvados). The exact provenance of some of the early finds (some of them already
known to Cuvier) is uncertain, and confusions with Callovian material from the
Vaches Noires have occurred.
The theropod material originally described as crocodilian by Cuvier in 1808
(Figure 2) from the collection of the Abbé Bachelet seems to have come mainly from
Early Kimmeridgian beds near Honfleur (Rioult, 1978a). It consists of vertebrae
now kept at the Paris Museum (see Taquet, 1984), which were figured as Streptospondylus cuvieri by Lennier (1870, 1887).
The Lower Kimmeridgian of the Cap de la Hève, on the north side of the Seine
estuary, has also yielded dinosaur remains. A theropod tooth found by Lennier was
described by Valenciennes (1863) and then made the type of the species Megalosaurus insignis by E. Eudes-Deslongchamps and G. Lennier (in Lennier, 1870). As
pointed out by Powell (1987), this species is based on undiagnostic material and
should be considered as invalid. The above-mentioned tooth was figured by Lennier
(1887) together with a few poorly identifred bones also referred to Megalosaurus
(Figure 4). A vertebra found by Lesueur at the Cap de la Hève was figured as
Srreptospondylus cuvieri by Lennier in 1870 (pl. VIII, fig. 2) and 1887 (pl. XIV, fig.
2); it was referred to a small sauropod by Huene (1926). All this material was kept in
the Natural History Museum at Le Havre and was destroyed by an American air raid
in September 1944.
In the Upper Kimmeridgian (Aulacostephanus mutabilis zone) of the cliffs of
Octeville (Seine-Maritime) , a few kilometres north of Le Havre, a partial skeleton of
a stegosaur was found by Savalle and Lennier in 1896 (Lennier, 1899). Despite
attempts by Gaudry to obtain the specimen for the Paris Museum, it was kept in Le
Havre, where it was studied by the Hungarian palaeontologist F. Nopcsa a few years
later (Nopcsa, 1911). The incomplete skeleton, consisting of vertebrae, elements of
the pelvis and a fragmentary femur, was made the type of a new species of
Omosaurus Owen, O. lennieri. As noted by Lucas (\902), however, Omosaurus
Owen, 1875 is preoccupied by Omosaurus Leidy,1856 (a phytosaur), and the
replacement name Dacentrurushas been suggested by Lucas. The type of Dacentrurus lennieri was destroyed with most of the palaeontological collections in Le Havre
by the air raid of September 1944.
Much more recently, the distal half of a stegosaur femur was found in the Upper
Kimmeridgian of the Octeville cliffs, about 1 km from the site of the discovery of the
type of Dacentrurus lennieri.It has been described by Galton and Boiné (1980) and
referred by them to Dacentrurw lennieri.
In 1887, Lennier referred to Tapinocephalus Owen a large vertebra and ribs from
the Kimmeridgian of the Cap de la Hève, which he considered as belonging to a
dinosaur. Tapinocephalr.rs is in fact a dinocephalian therapsid from the Permian of
South Africa, and Lennier's figure suggests a large sauropterygian. To add to the
confusion, the figure caption indicates "Tapinosauras." In 1923, Rabeck referred to
Tapinosaurus vertebrae and ribs from the Kimmeridgian of Octeville; to judge from
the photographs he published, they clearly belonged to a sauropterygian.
Tantalizing remarks about Kimmeridgian dinosaurs from the Le Havre area were
published by Marsh in 1897, after a visit to the museums in Caen and Le Havre.
Among undetermined material kept in Caen, Marsh "was greatly pleased to find the
2l
FRENCH DINOSAURS
t1v,1lrl"itl-,-lt:_1a
9.v:ii:ai {.ilt/,*;itL'.t* tlt:
7"*?m:tnô'a:"
-- 3'r'nzt
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11
Figure 4 " Megalosaurus" remains liom the Upper Jurassic of the Le Havre area, figured by Lennier in
1887. Although the tooth (1-3) does indeed belong to a theropod, the foot bones (4 8) are of more
doubtful attribution (the large ungual phalanx, 6-7. more probably belonged to a sauropod). Figure 9 is a
''section through a dinosaur bone," and fig. 10 is supposeclly a "bony scute from the skin of a
Me7alosaurus"t The littlc reconstruction at the top is meant to show Megalosaurus insignis. It is
reminiscent of the Crystal Palace model by Owen and Waterhouse Hawkins and was already much
outdated bv 1887.
28
E. BUFFETAUT, G. CUNYANDJ, LELOEUFF
geîus Pleurocoelus well represented by characteristic fossils, and from a well-defined
Jurassic horizonin the vicinity of Havre. " He thought this material belonged to a new
species, smaller than Pleurocoelus suffosus from the Kimmeridgian -of Swindon
(England), and closely resembling Pleurocoelus nanus , which he haâ described from
the Potomac Formation of Maryland (a resemblance he used to suggest that the
Potomac Formation was Jurassic in age-rvhereas it is now knowÀ to be Early
Cretaceous). In Le Havre, Marsh "found the bones of another Dinosaur, also one of
the Sauropoda, but considerably larger than rhe pleurocoelus at caen." The
remains, which came from the Kimmeridgian, were said to be "very similar to those
of Morosaurus." The material mentioned by Marsh was neithér described nor
!8ured, and was destroyed in 1944 with the rest of the palaeontological collections in
Caen and Le Havre. The same fate befell a scapulo-coiacoid from ihe Kimmeridsian
of_octeville, kept in Le Havre, and referred to pelorosaurar sp. by Huene
\l9rg).
Dinosaur remains have also been found in the Kimmerideian of-the Boulonnais.
near Boulogne-sur-Mer (Pas-de-Calais), although most oithe dinosaur materiai
from that area comes from the Portlandian. The dinosaurs from the Boulonnais have
been described by H. E. sau.vage in a number of papers between 1873 and 1914 (see
list in vadet and Rose, 1986). In his last paper on rhe subjecr, Sauvage (1914) li;ted
the following forms from the Kimmeridgian of the Boulonnais:-the supposed
sauropod Morinosaurus typul and an indeterminate sauropod, Megalosauris-insignis, Omosaurus and lguanodon sp. All of these identificaiions weie based on vely
scanty material. Morinosaurus typus was based on an isolated tooth of doubtful
affinities.
Two vertebrae of a "large saurian" from Alligny, near cosne (Nièvre), were
figured and described as possibly belonging to a megalosaurid by Gervais (1g59),
who mentioned that they may also have belonged to a plesiosaur. They are tiàteO âi
Megalosaurus sp. and attributed to the Kimmeridgianby Lapparent (uoz;. How-
ever, Gervais's figures do suggest sauropte{ygian rather than dinosaurian affinities.
Rather enigmatic trackways discovered in the course of systematic excavations in
th^e Late_Kim_meridgian lithographic limestones of Cerin (Ain, near the southern tip
of the Jura Mountains) have been attributed to leaping dinosaurs and nameà
Saltosauropus latus by Bernier et al. (1984). Further disèoveries have cast some
doubt on this interpretation, and the tracks in question may in fact have been left by
swimming animals pushing themselves with their hind feet on the bottom of a shallow
lagoon.
Portlandian As mentioned above, most of the dinosaur remains from the Boulon-
nais come from freshwater or fluviomarine beds referred to the Late Portlandian (or
sometimes to the Purbeckian by Sauvage). As early as 1839, a large bone from the
Portlandian was exhibited by Constant Prévost during a visit of thé Société géologi-
que de France; in 1880, the specimen was identified as the distal end of a fèmur 6v
Seeley, and referred by him to a cetiosaur. Most of the usually fragmentary dinosaur
remains from the Portlandian of the Boulonnais were described by Sauvage in a
series of notes, in which various and sometimes contradictory identifications were
gomelimes given to the same specimens. In his last paper on the subject, sauvage
(1914) listed the following forms: the sauropod Pelàrosaurus humerocristatus, [te
theropods Megalosaurus insignis and Megalosaurus oweni (distinguished by their
teeth), and the ornithopod Cumnoria all. prestwiclzi. In previous pàpers (see vadet
and Rose, 1986), the large spatulate teeth finally identified as Pàlorosiauras had
variously been referred by Sauvage to lguanodon praecursor Sauvage and to
Caulodon Cope, as well asto Neosodonby de la Moussaye (188s). Asto Cumnoria
FRENCH
DINOSAURS
29
aff. prestiwichi,il"had been identified on the basis of an isolated caudal vertebra,
described and figured as lguanodon prestwichi by Sauvage in 1897-98 (in a memoir
on fossil vertebrates from Portugal!). The following taxonomic history of this
particular fossil is extremely intricate (Rose, 1987) , since it was subsequently used as
type of the genus Teinurosaunzs by Nopcs a (1928, 1929) , and as type of the species
Caudocoelus sauvagei by Huene (7932), both considering it as belonging to a
theropod. Olshevsky (1978) has suggested that the name Teinurosaurus sauvagei
(Huene, 1932) should be applied to this vertebra. Whether it is distinctive enough to
warrant the erection of a taxon is very uncertain, however. Contrary to a widespread
opinion (expressed, for instance, by Lapparent",7967), the vertebra in question has
survived two world wars and years of neglect, like a large part of the other fossil
reptile remains in the collections of the Boulogne Natural History Museum (see
VadetandRose, 1986). Arevisionof theLateJurassicdinosaursof theBoulonnaisis
planned by the senior author. What can be said at the moment is that both a large
theropod and a large sauropod with spatulate, brachiosaurid-like teeth occur, the
latter being usually referred to the poorly known gents Pelorosaurus Mantell.
Fragments of very large bones in the Boulogne Museum indicate a sauropod of
enormous size.
In addition, washing and screening of sediment from two sites in the Upper
Portlandian of the Boulonnais by members of our department has resulted in the
discovery of a few dinosaur teeth indicating a theropod, a sauropod (possibly
Pelorosauru.s), a possible iguanodontid and what may be either a fabrosaurid or a
nodosaurid (Cuny et a|.,1991)
Little is known of the Portlandian dinosaurs of eastern France. In 1872. Pictet
briefly mentioned a few caudal vertebrae from the shallow marine limestones of the
Cyprina brongniarti zone of Ville-en-Blaisois (Haute-Marne), which he thought
were reminiscent of Megalosauras. These vertebrae are kept at the University of
Paris and have recently been revised by t\e senior author (Buffetaut, 1990), who
identified them as those of a sauropod (possibly a camarasaurid or a brachiosaurid).
The most complete dinosaur skeleton hitherto found in France was discovered in a
lithographic limestone quarry on the Petit Plan de Canjuers, a vast limestone plateau
north-west of Draguignan (Var), in Provence. It was described as a new species of
Compsognathus, C. corallestris, by Bidar et al. in 1972. Although the Canjuers
lithographic limestones have been referred to the Berriasian, notably by Fabre
(1981), the available evidence suggests that the original assignment to the Portlandian is correct. The skeleton, preserved on a slab of limestone, is generally well
preserved and nearly complete (the hands are very incompletely preserved, however). Bidar et al. (1972) had reconstructed the poorly preserved forelimb of the
Canjuers Compsognathus as a flipper-like structure implying semi-aquatic life
habits, on the basis of "wrinkle-like" impressions on the limestone slab. These have
been shown to extend well beyond the forelimb region by Ostrom (1978), who rightly
considered the flipper interpretation as "poorly founded and highly improbable."
Ostrom (1978) has found the Canjuers specimen to be nearly identical with the type
of Compsognathus longipes from the Solnhofen lithographic limestones of Bavaria
(although the French specimen, with a presacral column length
about 50% larger than the German one), and has thus referred
of
342 mm, is
it to the same
soecies.
A single theropod footprint has been described (as that of a coelurosaur) from the
Portlandian of Chassiron (on Oléron Island, Charente-Maritirne) by Lapparent and
Oulmi (1964). They also briefly mention a record of a theropod femur from the same
resion. We have found no additional information concernins the latter find.
E, BUFFETAUT, G. CUNY AND J. LE LOEUFF
6. Early Cretaceous
unexpectedly-enough, the French "wealden" of the Boulonnais and the pays de
Bray has yielded very few vertebrate remains, contrary to its English and Beigian
equivalents. Most of the Early Cretaceous dinosaurs from France hâve been founî itt
shallow marine rocks in the eastern part of the paris Basin, although there are
scattered records from other regions.
"wealden" The collection of the Lille Natural History Museum contains a theropod
claw and a large, possibly pathological vertebra of doubtful affinities, both fromihe
Wealden of Wimereux (Pas-de-Calais).
Valanginian Fore limb elements of a theropod have been found in the Valanginian
of crespian (Gard, southern France; M. Jeantet, personal communication).Hauterivian Some of the
best Early Cretaceous dinosaur material from France was
found during the last century near the town of Saint-Dizier (Haute-Marne), in the
eastern part of the Paris Basin, by J. Cornuel. This material was neglected for many
years, until Lapparent and Stchepinsky published a short note on it in 1968. Thê
dinosaur remains in the Saint-Dizier museum are currently being prepared and
revised by v. Martin at the university of Paris. The dinosaur remains in this
collection range in age from Hauterivian to Early Aptian.
The best material collected by Cornuel came from a quarry in the Hauterivian
shallow marine Calcaire à spatangues near wassy (Haute-Marne). Cornuel (1950)
mixed the dinosaur bones with remains of ichthyosaurs and plesiosaurs, and briefly
described the whole artificial assemblage under the aptly choien name oT Heterosoirus neocomiensis. Lapparent and Stchepinsky (1968) identified the dinosaur bones as
a partial lguanodon skeleton, comprising many vertebrae, part of the pelvis and limb
bones. They considered this animal as ve[y reminiscent of lguanodon mantelli.
Barremian The Lower Barremian of Baudonvilliers (Meuse)
has yielded a small rib
fragment (now in the saint-Dizier Museum) referred by Lappareni and Stchepinsky
(1968) to a small lguanodon.
The Paris Natural History Museum contains a large humerus from the non-marine
9pp". Barremian of Ville-sur-Saulx (Meuse, eastern part of the Paris Basin), which
Lapparent and Stchepinsky (1968) have referred to lguanodon bernissartensis.
Other Late Barremian lguanodon bones from the iame area include limb bone
fragments from oolitic ironstone near Saint-Dizier (Lapparent and Stchepinsky,
1968). A few bones of especially small size from the nôn-marine clayey sânos ôt
cousancelles (Meuse) were considered as those of a very young lguànodon by
Lapparent and Stchepinsky (1968).
rnl97l, various lguanodon remains, including a fairly complete skeleton, were
found when excavating a canal in Late Barremian non-marine deposits at SaintDizier (Damotte et a|.,1978). This material is still undescribed.
aptian. The Early Aptian marine clays of La Grange
au Ru, near wassy (HauteMarne), have yielded a large cubitus and a probable tibia fragment, nôw in the
Saint-Dizier Museum. Lapparent and Stchepinsky (1968) referred them to lguonodon bernissartensis.
Albian A
series of ten platycoelous sauropod caudal vertebrae was described by
Lapparent (1946) from the Albian of Villers-Saint-Barthélémy (Oise ), in the Pavs dê
FRENCH
DINOSAURS
31
Bray. A very similar isolated sauropod caudal vertebra from the Albian of Bléville
(Cap de la Hève, Seine-Maritime) has been found by Buffetaut (1984) in the
collections of the Rouen Natural History Museum.
Various dinosaur remains were described in the last century from the Albian of the
north-eastern part of the Paris Basin. The first report was by Barrois (i875), who
mentioned theropod teeth and vertebrae of Megalosaurus from the Gault of
Grandpré (Ardennes) and Louppy (Meuse), and a scute of Hylaeosaurus (an
ankylosaur) from Grandpré (that specimen, still in the collections of the Lille
Natural History Museum, is extremely doubtful). In 1876, Sauvage figured teeth and
vertebrae from Grandpré and Louppy, which he also referred lo Megalosaurus,
together with a limb bone fragment from the Gault of Boulonnais, which he thought
was reminiscent of Cope's Cionodon (a hadrosaur genus of doubtful validity). More
fossils, collected in the Albian phosphate-bearing beds of Louppy by Louis Pierson,
were described by Sauvage in 1882, together with additional material from Grand-
pré. The material from Louppy included a partial skeleton of a middle-sized
theropod (Figure 5), comprising teeth, vertebrae, ribs, a femur, part of a tibia and
several elements of a hand. Sauvage included the theropod material from the Gault
of eastern France in a new species of Megalosaurus, M. superbus. In 1926, Huene
remarked that the postcranial bones (especialy the femur) were very different from
those of Megalosaurus, and he erected the new genus Erectopus for the material
described by Sauvage. In 1932, he separated the teeth, which he thought were too
large to be associated with the bones and designated as "Gen. indeterm. sLrperbus,"
from the bones, which he called Erectopus sauvagei (Huene apparently overlooking
the fact that, according to Sauvage, Pierson had found some of the teeth in the same
block as the partial skeleton) . The limb bones of Ere ctopus are indeed different from
those of most other theropods (in 1932, Huene even placed Erectopus in a family of
its own, the Erectopodidae), and a revision would be welcome. Unfortunately, the
Pierson collection seems to have disappealed (even Huene apparently based his
observations only on Sauvage's description and ûgures).
In southern France, the Albian Grès verts of the Mont Ventoux, near Bédoin
(Vaucluse) have yielded a sauropod humerus, called Aepisaurus elephantinus by
Gervais (1848-52), who also described a few teeth from the same locality as possibly
belonging to the same form; however, the description suggests crocodilian rather
than dinosaurian affinities.
8. Late Cretaceous
Although Cenomanian deposits have yielded dinosaur remains in several parts of
France, little is known from the Upper Cretaceous of France prior to the Campanian.
On the other hand, the richest and most numerous dinosaur localities hitherto found
in France are in the Late Campanian to Maastrichtian continental formations of
southern France.
Cenomanian A few fragmentary remains from the Cenomanian Chalk of Normandy have been referred to dinosaurs (see Buffetaut, 1979). They include bones
from the "Craie chloritée" of Coulonges-sur-Sarthe (Orne) referred by Morière
(1871) to Iguanodoru and later interpreted by Bigot (1904) as dermal spines reminiscent of those of Stegosaurus, and a possible dermal scute of a dinosaur found by
Stiegelmann in the Cenomanian of the Cap de la Hève (Catherine, 1931). None of
this material was described or illustrated, and all the specimens were destroyed by
Allied bombings in7944. The systematic affinities and even the dinosaurian nature
of these remains are doubtful.
E. BUFFETAUT. G. CUNY AND J. LE LOEUFF
.,t
h
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of the most intriguing dinosaur specimens from France, described
superbus by Sauvage
as Megalosaurus
in 1887. The peculiar characters of this set of postcranial bones from the Albian of
Louppy (Meuse) led von Huene to consider them as belonging to a distinct genus and species, which he
called Erectopus sauvagei and placed in a family of their own, the Erectopodidae. The present
whereabouts of these interesting specimens are unknown, and all that is left are Sauvage's descriptions
and illustrations.
FRENCH
DINOSAURS
33
The palaeontological collection of the University of Le Mans contains a caudal
vertebra of a titanosaurid sauropod from the Cenomanian of Le Mans (Sarthe),
described by Buffetaut (1989).
The Cenomanian lignites of various parts of southern France have yielded few
dinosaur remains. Although Arnaud (1866) mentioned the occurrence of remains of
"gigantic saurians" in the lignites of the Sarlat region (Dordogne), this material was
never described and seems to be lost. A single theropod tooth from the lignites of
Saint-Julien-en-Peyrolas (Gard) was mentioned by Gervais (1859) and described
and figured as Megalosaurus by Dumas (1876); it also seems to be lost.
A humerus of a sauropod from the Cenomanian of Mondragon (Vaucluse) was
briefly mentioned by Depéret (1894) as resembling Aepisaurus, a conclusion disputed by Boule (1894). The present whereabouts of the specimen are unknown.
Several large dinosaur bones found when ploughing a field at Saint-Agnant, near
Brouage (Charente-Maritime), were briefly described as Megalosauras by Boisselier
(1882). The material, which includes amphicoelous caudal vertebrae and limb bone
fragments has recently been reexamined by two of us (E. B. and J. L. L.) at the La
Rochelle Natural History Museum, and found to belong to a sauropod. Its age is
somewhat uncertain: it comes from poorly dated beds overlain by Cenomanian
deposits. Although the dinosaur-bearing beds have sometimes been thought to be
terminal Jurassic in age, they are usually considered as basal Cenomanian.
Recently, washing and screening of Cenomanian sands at Lussant (CharenteMaritime) has resulted in the discovery of a serrated theropod tooth fragment and
what may be a worn tooth of a very small sauropod, together with more abundant
crocodilian teeth. This material is being studied by E. Buffetaut and D. Pouit.
Turonian The Turonian of Châteauneuf (Vendée) has yielded two very worn small
compressed teeth, which may belong to theropods. They will be described by E.
Buffetaut and D. Pouit.
Senonian Among reptile material from the Senonian of Notre-Dame-de-Riez
(Vendée) are three incomplete theropod teeth, to be described by E. Buffetaut and
D. Pouit.
Campanian Several of the important Late Cretaceous dinosaur localities of
southern France are of somewhat uncertain age, and although they are usually
referred to the Maastrichtian, some of them may in fact be Campanian in age (see
discussion in Buffetaut and Le Loeuff, in press). The finds listed below are from
relatively well dated Campanian localities, often in marine deposits.
As early as 1848-52, Gervais described the distal part of a sauropod humerus and
another limb bone fragment, from the "Craie tufau" of the Plaine de Lisle near
Périgueux (Dordogne). The age of this material is probably Campanian.
Much more recently, Platel (1989) has mentioned the discovery of four caudal
vertebrae of a titanosaurid sauropod in the marine Upper Campanian of the Ribérac
region (Dordogne).
Although remains of freshwater reptiles such as turtles and crocodilians are
relatively frequent in the lignite-bearing "Fuvelian" beds of Provence, very few
dinosaur bones have been reported. The only known specimens are two incomplete
vertebrae and the proximal part of a femur from a Fuvelian limestone at Le Beausset
(Bouches-du-Rhône), kept at the University of Lyon. They will be described as a
new genus and species of abelisaurid theropod, Tarascosaurus salluvicus, by Le
Loeuff and Buffetaut (in press).
34
E. BUFFETAUT, G. CUNY AND J. LE LOEUFF
Maastrichtian An incomplete hadrosaur dentary was described from the "Calcaire
Nankin" of Saint-Martory (Haute-Garonne) by paris and raquet in 1973. The
Calcaire Nankin is a shallow marine, possibly estuarine limestonè of probably Late
Maastrichtian age (Lepicard, 1985). Paris and raquet (1973) havô stressèd rhe
resemblances betwen the jaw fragment from Saint-Martory and material of Telmatosaurus transsylvanicas from Transylvania, but according to Brinkmann (19gg), the
specimen can only be considered as an indeterminate hadrosaurid.
Campanian to Maastrichtian We include under this rather imprecise stratigraphic
term the many dinosaur localities (numbering more than 80, including the eggshell
sites) in the Late Cretaceous non-marine deposits of southern Franèe, froir-Provence in the east to the Pyrenees in the west. Correlating these deposits with the
marine stratigraphic scale has proved difficult (see revièw in Bufietaut and Le
Loeuff, in press), although recent magnetostratigraphic work has provided somewhat better data, at least in Provence (Westphal and Durand, 1990). It seems that
most of the localities are Maastrichtian in age (possibly mostly early Maastrichtian),
but a I ate campanian age is not excluded for some of them (Éuffetaut and Le
Loeuff, in press).
The localities of Provence, Languedoc and the pyrenees have yielded a fairly
homogeneous dinosaur fauna, although some of the "rare" forms'(such as theropods) are not known in all localities, and the proportions of the different faunal
components vary from site to site.
At many localities, the assemblage is dominated by titanosaurid sauropods,
probably belonging to at least two distinct species, one of them being Hypselosàurus
priscus Matheron, 1869; although this taxon is often mentioned, it is poôrly defined
(Le Loeuff et al., 1989). The theropods are usually not abundani. Mentions of
Megalosaurus in the older literature are unfounded, being based on very scanty
material. In recent years, new theropod material has come to light, including smail
dromaeosaurid-like teeth (Buffe tavr et al . , 1986) from several localities in Provence
and Languedoc, and an abelisaurid maxilla from Provence (Buffetaut et al., 1988).
Material currently under study by two of us (J. L. L. and E. B.) includes dromaeosaurid postcranial remains from Provence.
, Ornithopods are mainly represented by Rhabdodon priscus, originally described
from Provence (Figure 6) by Matheron (1869), which has often been considered as an
iguanodontid, but should probably be placed among the Hypsilophodontidae (see
Brinkmann, 1988). Fladrosaurid remains are very rare and fragmentary.
Ankylosaur remains are not frequently found, and most of the French material
consists of armour plates, usually referred to Struthiosaurus Bunzel, 7871, a nodosaurid originally described from the Campanian of Austria, although Nopcsa (1929)
erected the taxon Rhodanosaurus lugdunensls for the French material.
In Provence, remains of a "large saurian" were reported as early as 1842 by
Doublier, from what remains one of the major French localities at Fox-Amphoui
(Var). Fairly abundant material has been collected there since that time. notablv bv
Lapparent, who described his finds in his 1947 monograph. Additional remains werê
collected in 1979-80 (Broin et a1.,1980), but no detailed description has been
published.
The first real description of dinosaur bones from the upper Cretaceous of
Provence was provided by Matheron in a paper published in 1869, in which he
described and named Hypselosaurtu priscus (which he still considered as possibly
allied to crocodiles) and Rhabdodon "priscum" (see Brinkmann, 1987, about thê
validity and proper gender of Rhabdodon), both from the "Rognacian" (a local
FRENCH DINOSAURS
35
*b;*tt.I*rlttv 1it'ixt ttttt :tra*i{'1,t * u lan,/b
:
1
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Figure 6 Partof thetypcmaterialof theornithopodR/raôdodonpriscus (dentaries,presumablyfromthe
same indiviclual), from ia Nerthe near Marseille, as illustrated by Matheron in the 1u69 paper in which he
lirst clearly identified dinosaurs in the continental Upper Cretaceous of southern France (although
remains of "large saurians" had previously been reported)
36
E. BUFFETAUT. G. CUNY AND J. LE LOEUFF
continental equivalent of the Maastrichtian, see Westphal and Durand, 1990) of the
Marseille and Aix-en-Provence region (Bouches-du-finOne). Matheron also was the
first to report the discovery of eggshell fragments, which he tentatively referred
either to a gigantic bird or to Hypselosaurus, in the upper Cretaceous of the Aix
basin. Despite isolated papers on the subject (Gervais^, lg77;yan Straelen and
Denaeyer, 1923), it was not until the 1950s that the remarkably rich egg sites of
Provence received the attention they deserved, thanks largely to ihe worÈôf Dughi
and Sirugue (see, for instance, Dughi and Sirugue, 195g, lgl6, ana. for more recJnt
work, Penner, 1985 and Jaeger, 1989). Although the eggs from provence have often
been.referred to Hypselosaurus, several disiinct typés are in fact present, but
identifications have hitherto been uncertain for IaôÈ of direct assoôiations with
skeletal material. Although a number of dinosaur bones have been found in the Aix
basin, no major site with skeletal remains is known in the Aix resion. with the
possible exception of Saint-Estève-Jeanson, where fairly abunclant-maierial was
found in the 1960s but has never been described.
Several localities (with bones or eggs) are known in Languedoc, west of the Rhône
valley, r.n th9 departments of Gard and Hérault. very little has been publishecl
concerning the material
Jounf ll the regions around Nimes (,,Champ-Garimond":
see Ledoux et a1., 1966) and Montpellier (including the viileveyrac basin, from
which "large saurians" were reported by Bleicher aslarly as rgTi). The dinôsaurs
found farther west, in several localities around Saint-Chinian (Hérault), are better
known. Although Gervais mentioned the occurrence of forms resemblins those from
Provence in this area as early as 1877 , and remains of large reptiles weràmentioned
by Miquel in 1895, the first important finds were reporteà by bepéret at the turn of
the century.(pgRéret,.1899, 1900a, 1900b), and more maieriai was described by
Lapparent (1941 ,7954). Some of those localities still yield bones, and a fairly weil
preserved Rfr abdodon dentary, ind_icating a new species, was found at one of thêm by
two of us in 1990 (Buffetaut and Le Loeuff, in press).
still farther west, in the corbières, both eggs and bones have also been found. Egg
localities were the first
to attract the attention of palaeontologists, notably îi
Saint-André-de-Roquelongue and Rennes-le-Château (Aude). Thè nestins sités at
Rennes-le-Château have recently been described (cousin er al.. l9g9l. Àtthough
dinosaur bones were reported from the l-lpper Aude valley by Gervaii as earlyîs
1877, the importance of the localities in this area was not recôsnized until 1983. when
Clottes and Raynaud described specimens from several néwly discovered sites.
Systematic excavations conducted by our group at several localitËs around the town
of Espéraza(Aude) in 1989 and 1990 have yielded an abunclance of dinosaur bones,
among which titanosaurid remains are especially abundant. The sites of the Uppei
Aude Va-tley are clearly among the richest dinosaur localities in the Upper Ôietaceous of France (Buffetaut et al., 1989)
The foothills of the Pyrenees, in department Ariège, have also yielded dinosaur
remains. A chance find ofsauropod and Rhabdodon bones in the Upper Cretaceous
of Lavelanet was described by Villatte et al. (1986). Much eailier. vertebrate
remains. including those of dinosaurs, were found around the town oI Le Mas d'Azil
by Abbé J. J. Pouech, a local priest, who first recorded these finds in 1859. Althoush
he later wrote to Gaudry about an especially rich locality (pouech, lggl), pouecË's
discoveries went unnoticed. His important collection has recently been rediscovered in a religious school in Pamiers (Ariège) by one of us (J. L. L. ); it includes
remains of titanosaurids. nodosaurids and Rhabdodon (Le Loeuff. in press). pouech
also was the first to describe dinosaur eggshells, although the exact nature of his finds
was not fully clear to him (Buffetaut and Le Loeuff, 1989). Todav, most of pouech's
FRENCH
DINOSAURS
31
localities are completely overgrown and no longer accessible, although we have been
able to find eggshell fragments at one of his sites.
REWORKED DINOSAUR REMAINS
A few reworked dinosaur teeth have been found in Miocene deposits of the Loire
Valley. An iguanodontid tooth from the Late Miocene "Faluns" of Doué-laFontaine (Maine-et-Loire) was described by Buffetaut et al. (1980), and two small
theropod teeth from the same deposits have been referred 1o Carcharodontosaurus
saharicus (Depéret and Savornin, 1927) by Mornand (1989); the latter teeth are
better considered as belonging to an indeterminate theropod. All these teeth were
probably reworked from underlying Cenomanian beds.
DINOSAUR REMAINS OF UNKNOWN PROVENANCE
In 1870, Gaudry figured and briefly described the distal end of a dinosaur femur given
to the Paris Museum by the "Petit Séminaire" in Beauvais (Oise). The origin of the
specimen is unknown. àlthougtr Lapparent (1967)listed it as coming from the Lower
Cretaceous of the Pays de Bray. Gaudry only identified it as dinosaurian. It appears
as an ornithopod in Lapparent's list (1967), but its morphology rather suggests a
theropod.
CONCLUSIONS
The most obvious result of this brief revie',v is to show that dinosaur remains are
known from many geological horizons in many parts of France. As in other parts of
the world, the completeness of the record seems to be partly dependent on its age,
and this in turn is to some extent linked to changing depositional environments,
themselves conditioned by global sea-level changes (see Haubold, 1990). The French
record is thus relatively good for periods of widespread emersion, in the Late
Triassic, Late Jurassic and, particularly, Late Cretaceous. The relatively poor Early
Cretaceous record is somewhat surprising, especially by comparison with what is
known from that period in England and Belgium. All in all, however, the French
record is especially complete from a stratigraphic point of view, and in this particular
regard it is better than that of any other European country.
Although a large part of the dinosaur remains from France are fragmentary, some
are sufficiently complete to contribute to our knowledge of the anatomy of various
forms, and consequently to reconstructions of dinosaur phylogeny. This applies, for
instance, to some of the prosauropod material from eastern France, to the Callovian
stegosaur from Normandy, to the Oxfordian dinosaur from Damparis, and to the
Compsognathas specimen from the Portlandian of Canjuers. An important aspect of
some of the finds is that, coming from marine deposits, they are particularly well
dated: the Middle and Late Jurassic specimens from Normandy and the Early
Jurassic lguanodon remains from eastern France are good examples. The importance of other finds probably lies more in their biogeographical significance: the
dinosaur assemblages from southern France provide interesting evidence about the
Gondwanan affinities of many elements of the Late Cretaceous terrestrial faunas of
Europe (Buffetaut, 1989; Le Loeuff, in press). The numerous eggshell localities in
the Upper Cretaceous of southern France, which, if properly studied, can provide
imnortant evidence on dinosaur reproduction. should also be mentioned.
Èinally, it should be pointed oui that many gaps in our knowledge of French
E. BUFFETAUT, G. CUNY AND J. LE LOEUFF
38
dinosaurs are certainly due to insufficient interest in them on the part of French
palaeontologists, especially during the first half
of the twentieth century. Our
experience of the last few years clearly shows both that a number of specimens are in
need of a thorough revision, and that many important localities deserve a systematic
exploitation they have never undergone.
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