Eric Buffetaut. Encyclopedia of Dinosaurs. Editor: Philip J Currie & Kevin Padian. Amsterdam: Academic Press, 1997.
The first dinosaur remains to be described scientifically were found in Europe at the beginning of the 19th century. Since then, however, European dinosaurs, which are often represented by incomplete specimens, have been somewhat overshadowed by finds of spectacular complete skeletons from other parts of the world, including North America, Africa, and Asia. Nevertheless, the European dinosaur record, which includes skeletal remains, footprints, and eggs, is probably the most complete in the world from a stratigraphic point of view, with relatively few gaps in a series of sites covering the time span from the Late Triassic to the end of the Cretaceous.
Many European countries have yielded remains belonging to various groups of dinosaurs and a stratigraphic rather than systematic or geographic presentation seems to be the most appropriate way to present the European record in a concise way. Only the main sites and assemblages have been included.
In central and western Europe, the Late Triassic is largely represented by continental deposits, in which dinosaur bones and footprints sometimes occur in great abundance. Although there have been reports of dinosaur footprints in rocks older than the Late Triassic in several parts of Europe, including England and France, they appear to be erroneous or inconclusive. The European dinosaur record actually begins with small three-toed footprints from the Carnian of northern Bavaria and possibly from the Ladinian–Carnian boundary in the Swiss Alps. The earliest skeletal remains are Norian in age. The earliest relatively well-known European dinosaur is the prosauropod Sellosaurus gracilis, from the lower Stubensandstein of Württemberg, considered early Norian in age. Slightly more recent, probably middle Norian, levels of the Stubensandstein of the same region have yielded remains of the theropod Halticosaurus. Dinosaur bones first become abundant, however, in higher levels of the Late Triassic usually referred to the late Norian. Sites of that age include the famous German prosauropod localities in the Knollenmergel, such as Trossingen in Württemberg and Halberstadt in Sachsen-Anhalt, which have yielded many well-preserved skeletons of Plateosaurus, and various other sites in other parts of Germany, including the Grosse Gleichberg in Thuringia, where the theropod Liliensternus has been found. Outside Germany, Plateosaurus remains have been found in abundance in the upper Norian of Switzerland and in rocks of the same age in the French Jura mountains. Late Triassic dinosaur remains from Britain include the prosauropod Thecodontosaurus from the Magnesian Conglomerate near Bristol, which is Norian to Rhaetian in age. Dinosaur remains from the controversial “Rhaetian” stage, at the top of the Triassic, in various European countries (Britain, France, Germany, and Belgium) are usually very fragmentary elements from bone beds that are not easily identifiable; some can be referred to indeterminate prosauropods. An exception is the melanorosaurid prosauropod Camelotia from Somerset, England.
Norian dinosaur footprints, mostly attributable to theropods, are known from several sites in northern Bavaria, and also from northern Italy. Several footprint sites of Rhaetian age are known from France, the most important one being at Le Veillon (Vendée), on the Atlantic coast. There, hundreds of footprints have been found, most of them referable to theropods (Grallator and Eubrontes), although some have been attributed to ornithischians. “Rhaeto-Liassic” tridactyl footprints have also been reported from Scania in southern Sweden.
In Europe, the Early Jurassic is represented largely by marine deposits in which dinosaur remains are relatively infrequent. An incomplete theropod skeleton, recently redescribed as Liliensternus airelensis, comes from beds very close to the Triassic–Jurassic boundary, probably basal Hettangian in age, at Airel in Normandy (northwestern France). In England, more or less complete skeletons of the early thyreophoran Scelidosaurus harrisonii have been found since the 19th century in the marine Sinemurian of the Dorset coast. Very incomplete remains of another possible thyreophoran,Lusitanosaurus, were reported from the Sinemurian of Portugal. A third primitive thyreophoran, Emausaurus ernsti, was described on the basis of a skull and some postcranial elements from the Toarcian of Mecklenburg-Vorpommern in northern Germany. Hindlimb elements of a sauropod from the Toarcian Posidonienschiefer of Baden-Württemberg (southwestern Germany) have been described as Ohmdenosaurus liasicus. Besides these scattered skeletal remains from marine deposits, which also include a few isolated theropod bones or teeth from England, Scotland, France, and Germany, dinosaur footprints of Early Jurassic age occur in several European countries, often in calcareous rocks deposited in a beach or mudflat environment. Such trackways, mostly from theropods, are known from the Hettangian and Pliensbachian of the southern rim of the French Central Massif. Other Early Jurassic dinosaur footprints, most of them referable to theropods, have been reported from the Hettangian of Hungary and the Holy Cross Mountains of Poland. Among the early Liassic footprints from the Rovereto area in northern Italy, some have been referred to a very early sauropod.
The Middle Jurassic dinosaurs of Europe are better known than those from the Lower Jurassic mainly because of a few comparatively rich localities in England and France. Bajocian forms are poorly known, although remains of the theropod Megalosaurus have been described from the Inferior Oolite of Dorset, England. Dinosaur remains have been found in abundance in several Bathonian formations in England, including the famous Stonesfield “Slate” which yielded the remains of Megalosaurus bucklandi, the first dinosaur to receive a proper generic name, in 1824. The Bathonian of England also contains remains of the large sauropod Cetiosaurus oxoniensis and stegosaurs. In France, the Bathonian of the Caen area in Normandy yielded remains of a large theropod, that was described as Poekilopleuron bucklandi, but may in fact belong to Megalosaurus. Theropod and sauropod teeth are also known from the marine Bathonian of Saint-Gaultier in central France.
The Callovian dinosaurs of Europe are relatively well known because of fairly numerous finds from the marine Lower Oxford Clay of England and its equivalents in Normandy. The English finds include theropods such as Eustreptospondylus oxoniensis, represented by a nearly complete skeleton, sauropods (Cetiosauriscus andOrnithopsis), ornithopods (Callovosaurus), stegosaurs (Lexovisaurus) and one of the oldest known ankylosaurs (Sarcolestes). The French record from Calvados in Normandy is somewhat less diverse but nevertheless includes theropods (with at least two forms, one of which seems to be referable to Megalosaurus; the status of Piveteausaurus, known from a braincase from Normandy, is uncertain), stegosaurs (with a partial skeleton of Lexovisaurus), and sauropods. Middle Jurassic dinosaur footprints have been reported from several sites in England.
The Late Jurassic European dinosaur record is good, with important localities in several countries. Oxfordian theropod remains are known from marine deposits in England (with Metriacanthosaurus parkeri from Dorset) and Normandy (with Megalosaurus bones from the Vaches Noires Cliffs of Calvados). Isolated theropod and sauropod teeth have been found in the Oxfordian fluvio-marine Sables de Glos of Normandy. One of the best sauropod skeletons ever found in Europe comes from the Oxfordian of Damparis (Jura, eastern France); it belongs to a brachiosaurid and was found associated with several theropod teeth, indicating in situ scavenging on the sauropod carcass during a phase of emersion of a carbonate platform. In Portugal, the Guimarota lignite mine, of Late Oxfordian or Kimmeridgian age, has yielded hypsilophodontid teeth and, recently, teeth referred to Archaeopteryx.
Kimmeridgian dinosaurs are known from the Kimmeridge clay of England. They include poorly known sauropods, stegosaurs (Dacentrurus), and ornithopods (including a fairly complete skeleton of Camptosaurus prestwichii from the vicinity of Oxford). The marine Kimmeridgian of the Normandy coast near Le Havre also contains dinosaur remains, including theropods, sauropods, and stegosaurs. In the Jura mountains of Switzerland, abundant sauropod tracks have been found in Kimmeridgian limestones, as well as an incomplete sauropod skeleton. Other sauropod footprints are known from rocks of the same age in Barkhausen in northern Germany. A few dinosaur bones have been reported from the marine Kimmeridgian of the Boulonnais, in northern France, but most of the remains from that area are from nonmarine Tithonian rocks. They include bones and teeth of a large theropod, a large camarasaurid sauropod (Neosodon), and a small iguanodontid. The small theropod Compsognathus is known from two fairly complete skeletons from Tithonian lithographic limestones, in northern Bavaria and southeastern France.
Abundant dinosaur remains have been discovered in the Upper Jurassic (Kimmeridgian and Portlandian) of Portugal. The Kimmeridgian assemblage includes several sauropods, including a probable brachiosaurid, theropods, stegosaurs (Dacentrurus), an early ankylosaur (Dracopelta), and ornithopods. Footprints (of theropods and sauropods) are frequent in the Portlandian. The discovery of numerous dinosaur eggs in the Kimmeridgian of the Lourinha region is especially noteworthy. In Asturias (northern Spain), large footprints referable to sauropods have been reported from Kimmeridgian rocks.
The most varied Early Cretaceous dinosaur assemblage in Europe comes from the non-marine Wealden beds of southern England (Sussex and Isle of Wight), which range in age from late Berriasian to early Aptian. The Wealden fauna includes several species of Iguanodon; other ornithopod genera such as Hypsilophodon, Valdosaurus, and Vectisaurus; the nodosaurids Hylaeosaurus and Polacanthus; the early pachycephalosaur Yaverlandia; several sauropods (including a brachiosaurid, a titanosaurid, and a possible diplodocid); and several theropods, among which is the probable spinosaurid Baryonyx. On the continent, the most famous Wealden dinosaur locality is undoubtedly the Bernissart coal mine in Belgium, where about 30 skeletons of Iguanodon, belonging to the species I. bernissartensis and I. atherfieldensis, were discovered in 1878. Dinosaurs also occur in the Wealden of northern Germany, where skeletal remains (including those of a theropod, an ornithopod, and the basal marginocephalian Stenopelix) are usually far less abundant than footprints. The latter include tracks of sauropods, ornithopods, and theropods from the region around Hanover (Bü ckeberg and Mü nchehagen). An exception is the Nehden locality in Westphalia, a karstic deposit of Aptian age, in which abundant remains of I. bernissartensis and I. atherfieldensis have been discovered.
In the eastern Paris Basin, a succession of alternating shallow marine and non-marine beds, ranging in age from Hauterivian to Aptian, has yielded well-dated Iguanodon remains that show a succession of species similar to that from the English Wealden. Remains of a brachiosaurid sauropod are also known from the Barremian of that area. In southern France, a few bones of an Allosaurus-like theropod have been described from the marine Valanginian of the department of Gard.
Early Cretaceous non-marine rocks in eastern Spain contain dinosaur assemblages. The mainly Barremian beds of the Galve area (Teruel Province) have yielded hypsilophodontids, theropods, and four forms of sauropods, including the camarasaurid Aragosaurus. From the early Aptian of the Morella region (Castellon Province), theropods, sauropods, ankylosaurs, and Iguanodon have been reported. Early Cretaceous ankylosaurs, iguanodontids, and theropods are also known from Burgos Province in north-central
Spain. In Cuenca Province in central Spain, the late Hauterivian to early Barremian lacustrine lithographic limestones at Las Hoyas have yielded a few dinosaur remains, including the early multitoothed ornithomimosaur Pelecanimimus. Abundant ornithopod and theropod footprints are known from the Lower Cretaceous of the La Rioja region in northern Spain. A few skeletal remains (of theropods, sauropods, and ornithopods) and tracks are also known from the Lower Cretaceous (Hauterivian and Aptian) of Portugal.
Among the few dinosaurs reported from Italy is the skeleton of a very small theropod from the Aptian lithographic limestones of Pietraroia (Benevento Province).
In eastern Europe, the most important Early Cretaceous dinosaur assemblage is that from the karstic bauxite deposits (Barremian to Aptian in age) of Cornet, in Transylvania (Romania). It mainly contains ornithopods (Iguanodon,Valdosaurus, and Dryosaurus), as well as a theropod.
On the northern outskirts of Europe, ornithopod and theropod footprints have been reported from the Barremian of Spitzbergen.
Albian dinosaurs are known from localities in England and France. The English material mainly comes from the Cambridge Greensand, mostly as disarticulated and reworked material. It includes sauropods, ornithopods (including a probable iguanodontid and a hadrosaurid), and ankylosaurs. In France, sauropod remains have been found in the Albian of the Pays de Caux and the Pays de Bray in Normandy, as well as in the “Gault” clay of the eastern Paris Basin. Dinosaurs from the phosphate-bearing Albian of northeastern France include the enigmatic theropod Erectopus. In southeastern France, a sauropod humerus was found in the Albian green sandstones of the Mont Ventoux.
The European dinosaurs of the early part of the Late Cretaceous, up to the Campanian, are relatively poorly known. In England, the nodosaurid ankylosaur Acanthopholis horridus comes from the Cenomanian Chalk Marl, and an ornithopod has been reported from the Totternhoe Stone, also Cenomanian in age. A few isolated and fragmentary specimens of sauropods and theropods are known from the Cenomanian, Turonian, and Santonian of west-central France. A few dinosaur remains (belonging to a theropod and to the iguanodontid Craspedodon) have been collected from the Santonian of Lonzée in Belgium. Cenomanian footprints are known from Portugal (theropods and sauropods) and from Croatia (sauropods).
The European dinosaur record becomes tolerably good again in the Campanian. A few isolated specimens are known from marine rocks of that age in southern Sweden (theropod and ornithopod) and southwestern France (sauropod). One of the best early Campanian localities was found in lignite-bearing beds at Muthmannsdorf in Austria in the 19th century; it yielded a theropod, the ornithopod Rhabdodon, and the nodosaurid ankylosaur Struthiosaurus.In southern France, the Villeveyrac locality, of the same age, also contains Rhabdodon, an ankylosaur, and a small theropod. In southeastern France, an abelisaurid theropod, Tarascosaurus, has been reported from Campanian beds at Le Beausset (Var).
Maastrichtian dinosaurs are known from marine Chalk deposits in the Limburg region of Belgium and the Netherlands, near the city of Maastricht. They include a theropod of uncertain affinities (Betasuchus) and hadrosaurs. Hadrosaur remains have also been found in the marine upper Maastrichtian of the Bavarian Alps, and there is one hadrosaur record from the marine Maastrichtian of the Crimea in Ukraine. However, most of the Maastrichtian dinosaurs from Europe have been found in non-marine rocks in Transylvania, in southern France, in northern Spain, and in Portugal (some of the French-Iberian localities may actually be late Campanian in age). The Transylvanian assemblage is apparently late Maastrichtian in age. It includes Rhabdodon, the primitive hadrosaur Telmatosaurus, the nodosaurid ankylosaur Struthiosaurus, titanosaurid sauropods (Magyarosaurus), and poorly known theropods. Also worth mentioning is the occurrence of eggs associated with remains of hadrosaur embryos.
Whereas the Portuguese record is poor (with indeterminate theropods and ornithischians), there are good late Campanian to Maastrichtian dinosaur localities in various parts of Spain. In the Tremp basin of Catalonia, sauropods, hadrosaurs, and Rhabdodon occur in late Maastrichtian deposits. Footprints (including those of sauropods) and eggs have also been found. Other Late Cretaceous dinosaurs are known
from various provinces of Spain. One of the richest sites is at Lano, near Vitoria in the Basque Country. It is of late Campanian to early Maastrichtian age and has yielded bones and teeth of abelisaurid theropods, titanosaurid sauropods, Rhabdodon, and the ankylosaur Struthiosaurus.
In southern France, dinosaur localities of early Maastrichtian age are known in great number in nonmarine formations extending from Provence in the east to the foothills of the Pyrenees in the west. The best known assemblages are from the Fox-Amphoux area in Provence and the upper valley of the Aude River. Skeletal remains indicate the occurrence of abelisaurid and dromaeosaurid theropods, armored titanosaurid sauropods, the ornithopod Rhabdodon, and an ankylosaur. Eggs are abundant in some areas, especially the Aix-en-Provence basin and the upper Aude valley. Although several types have been distinguished on the basis of shell microstructure, no clear associations with skeletal remains have yet been reported. Less numerous late Maastrichtian localities are also known, mainly from the Corbiéres region and the area of the Garonne valley. They contain a fauna dominated by hadrosaurs, accompanied by theropods and ankylosaurs. The southern French record thus suggests a faunal change, marked by a decline of titanosaurid sauropods and an expansion of hadrosaurs, during the Maastrichtian.
From this brief review, it appears that the European record covers most of dinosaur history in a remarkably complete way. In the Triassic and Jurassic, European dinosaur faunas showed clear resemblances to those of other continents, notably North America and Africa. In the Early Cretaceous, resemblances to North America were still marked (with such genera as Iguanodon and Polacanthus in common). In the Late Cretaceous, however, the European assemblages showed characteristics of their own, with taxa of “Gondwanan” affinities such as abelisaurid theropods and titanosaurid sauropods playing an important part together with endemic forms, whereas taxa of “Asiamerican” affinities were few. This of course reflects the changing paleogeographical history of the European continent during the Mesozoic, which alternatingly favored or limited faunal exchanges with North America, Asia, and Africa. The details of this complex biogeographical history still have to be worked out in detail.