Tetrapods from the Late Jurassic and Early Cretaceous lithographic limestones of Europe: A comparative review

TETRAPODS FROM THE LATE JURASSIC AND EARLY CRETACEOUS LITHOGRAPHIC

LIMESTONES OF EUROPE : A COMPARATIVE REVIEW

ERIC B U F F E T A U T Universit~ P. et M. Curie, Laboratoire de Pal~ontologie des Vertebras et URA 720 du CNRS, 4 place Jussieu, F-75252 Paris eedex.

BUFFETAUT E. Tetrapods from the Late Jurassic and Early Cretaceous lithographic limestones of Europe : a comparative review. [Les t~trapodes des calcaires lithographiques du Jurassique sup~rieur et du Cr~tac~ inf~rieur d'Europe : une revue comparative.] GEOBIOS, M.S. n ° 16 : 259-265.

ABSTRACT

Comparison of the tetrapod assemblages from Late Jurassic (Franconia, Cerin, Canjuers) and Early Cretaceous (Montsec, Las Hoyas) lithographic limestone localities in Germany, France and Spain, reveals both important ressemblances and striking differences. The latter are explainable by different collecting histories, taphonomic biases, different depositional environments, and differences in geological age. The tetrapod assemblages from the lithographic limestones of Europe are unbalanced and the reasons for this are worthy of further investigation.

KEY-WORDS : LITHOGRAPHIC LIMESTONES, TETRAPOD FAUNAS, LATE JURASSIC, EARLY CRETACEOUS, GERMA- NY, FRANCE, SPAIN, PALAEOECOLOGY.

RI~SUME

Une comparaison des assemblages de t~trapodes des calcaires lithographiques du Jurassique sup~rieur (Franco- nie, Cerin, Canjuers) et du Cr~tac~ inf~rieur (Montsec, Las Hoyas) d'Allemagne, de France et d'Espagne r~v~|e la fois des ressemblances importantes et des differences remarquables. Ces derniers s'expliquent par une histoire diff~rente des r~coltes, des biais taphonomiques, des milieux de d~p6t diff~rents, et des differences d'~ge g~ologi- que. Les assemblages de t~trapodes des calcaires lithographiques d'Europe sont d~sg~tuilibr~s, et les raisons de ce d~s~quilibre meritent des ~tudes plus pouss~es.

MOTS-CL]~S : CALCAIRES LITHOGRAPHIQUES, FAUNES DE T]~TRAPODES, JURASSIQUE Sb~PERIEUR, CRETAC]~ IN- FI~RIEUR, ALLEMAGNE, FRANCE, ESPAGNE, PAL]~OI~COLOGIE.

INTRODUCTION

Ever since the first studies on fossil reptiles from the lithographic limestones of Franconia at the end of the 18th century, the attention of palaeontolo- gists has been drawn to the often excellent preservation of te t rapod skeletons in fine grained rocks of tha t kind. Later, the discovery of such rarities as the A r c h a e o p t e r y x skeletons has added to the celebrity of lithographic limestones as a source of remarkable te t rapod remains. There is no question that the s tudy of the fossils from the various lithographic limestone localities in the Upper Jurassic and Lower Cretaceous of Europe has contributed greatly to our knowledge of the tetrapod faunas of these periods. It is also clear, however, that the tetrapod assemblages found in those limestones are peculiar in several ways. They certainly do not

correspond to "normal", balanced faunas, and, despite general ressemblances, there are noticeable differences in their composition from one site to another. The purpose of the present paper is to review briefly these assemblages, to compare them with each other, and to discuss the conclusions that can be drawn from such comparisons, especially from a palaeoecological point of view.

THE EVIDENCE

Tetrapod remains have been found in all the major lithographic limestone localities in Eu- rope ; the Altmthhl valley in northern Bavaria (Franconia), Nusplingen in Wiirttemberg, Cerin in the French Ju ra mountains (Department

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Ain), Canjuers in Provence (Department Var), Montsec (often written Montsech until recently) in Catalonia and Las Hoyas in Cuenca Province, Spain. The amount of information available about these various assemblages is extremely variable, because they have not all been studied to the same extent. The scientific literature about the Bavarian localities spans several centuries and is extremely abundant, whereas sites such as Canjuers and Las Hoyas have been discovered much more recently and have been the subject of much fewer studies.

The present paper is largely based on recent re- views of the faunas from the above-mentioned localities : Barthel (1978) and Barthel et al. (1990) for the Franconian sites, Barale et aL (1985) for Cerin, Fabre et al. (1982) for Canjuers, Barale et al. (1984) and Martinez-Delclos et al. (1991) for Montsec, and Sanz et al. (1988) for Las Hoyas. More detailed bibliographic lists are to be found in some of these papers. When needed, information from more recent or more specialized papers has also been used. No recent synthesis about the fauna from the lithographic limestones of Nus- plingen seems to be available, and this locality has therefore not been taken into consideration in the comparison, although remarks about some of the specimens found there have been made.

This comparative review is of course based on systematics, and this has entailed various problems. A major difficulty is that our knowledge of the various groups of tetrapods found in the lithographic limestones of Europe is quite varia- bles : some (e.g. pterosaurs) have been revised recently and their systematics are generally relia- ble, whereas others (e.g. ichthyosaurs) are still poorly known. In some instances, many systematic uncertainties remain. Sphenodonta are an interesting example : despite the revisions by Cocu- de-Michel (1963) and Fabre (1981), the classifica- tion of the Late Jurassic sphenodonts is still in a state of chaos, even at the family level. Because of this state of affairs, comparisons have not been attempted at low taxonomic levels. This review is based only on skeletal remains. Ichnological evidence has not been used, not because it is considered as unimportant, but because of interpretation problems (for instance, the tracks from Cerin described as those of hopping dinosaurs by Ber- nier et al., 1984, are interpreted as those of turtles swimming in shallow water by Thulborn, 1990), and also because undoubted tetrapod footprints have been reported only from Cerin (this is possibly due to the special excavation techniques used there during the last fii~een years, involving the uncovering of fairly large

surfaces along bedding planes, which is clearly favourable to the discovery of trackways).

THE A G E OF T H E L O C A L I T I E S

Dating some of the lithographic limestone localities of Europe has proved difficult. It is now clear that they are not all of roughly the same age, as was sometimes admitted, and faunal differences may be due at least in part to differences in geological age. The main "Plattenkalk" beds of the Altmfihl valley are considered as Tithonian in age (Barthel et al. 1990). The lithographic limestones of Cerin are placed in the Upper Kimmer- digian. The Spanish localities, Montsec and Las Hoyas, are younger in age, Montsec corresponding to the Berriasian-Valanginian transition (Barale et al. 1984), while Las Hoyas is referred to the Late Hauterivian-Early Barremian (Sanz et al. 1988).

The age of the Canjuers locality has been dispu- ted : after being placed in the Portlandian, it was long considered as Berriasian in age, an opinion still expressed in recent papers (Broin et al. 1991), Recent evidence based on crinoids indica- tes a Tithonian age (Roman 1991) a conclusion recently fully confirmed by ammonites (Atrops, this volume).

T H E T E T R A P O D S F R O M T H E LITHO- G R A P H I C L I M E S T O N E S : AN ANNOTA- TED R E C A P I T U L A T I O N

The list below is a succinct review of what is currently known of the tetrapods (amphibians, reptiles and birds) currently known from the Late Ju- rassic and Early Cretaceous limestones of Ger- many, France and Spain (see also table 1). For the reasons mentioned above, no effort has been made to reach a level of great systematic preci- sion. When needed, remarks have been made con- cerning the mode of life and ecological preferen- ces of certain forms.

CAUDATA

Several specimens of an indeterminate salaman- der were reported from Las Hoyas by Sanz et al. (1988). No remains of Caudata are known from the other lithographic limestone localities under discussion.

A N U R A

Three genera of frogs are known from Montsec (Montsechobatrachus, Neus ibatrachus and Eodis-

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"~ u) u'} Z ¢~ ~ o u=

~ o z z 7-

U u- L)

X CAUDATA

X ANURA

X X X X ICHELONIA

X ICHTHYOPTERYGIA

X SAUROPTERYGIA I,

X X X SPHENODONTA

X I X X SQUAMATA

X X X X X CROCODYLIA

X X X PTEROSAURIA

X X X DINOSAURIA

X X X AVES

Table 1 - The distr ibution of major groups of tetrapods in the Late Jurass ic and Ear ly Cretaceous li thographic l imestone localities of Europe (see text for de ta i l s about the palaeoecological significance of each group). Table based only on skeletal remains : ichnological evidence is not used. Distribution des grands groupes de tdtrapodes dans Ies gisements de calcaires lithographiques du Jurassique supdrieur et du Crgtacd infg- rieur en Europe (voir le texte pour des ddtails sur la significa- tion palgodcologique de chaque groupe). Tableau fond~ unique- ment sur los restes squelettiques :les donnges ichnologiques ne sont pas prises en compte.

have lived both in fresh water and in coastal environments.

ICHTHYOPTERYGIA

Ichthyosaurs are known only from the Franco- nian localities, where they are unfrequent. No recent revision is available. Barthel e t a l . (1990) mention two genera (Leptopterygius and Ma- cropterygius).

SAUROPTERYG~

The only sauropterygian remain from the lithographic limestones seems to be a very large plio- saur tooth from Franconia, referred to Stretosau- rus by Barthel etal. (1990).

SPHENODONTA

They are relatively abundant in Francoma, at Cerin and at Canjuers, but have not been reported from the Early Cretaceous Spanish sites (Meyasaurus from Montsec is a lacertilian, as shown by Hoffstetter in 1965). As mentioned above, these forms are poorly understood from a systematic point of view. Some were terrestrial (Ho- rnoeosaurus, Sapheosaurus), with possible digging habits in some cases, whereas others (Pleurosau- rus), with much elongated bodies were clearly adapted to life in the water.

coglossus : seeWenz 1991). Anurans are unknown from the other localities.

CHELONIA

Turtle remains, usually in the shape of fairly well preserved specimens, have been found in the Franconian localities, at Cerin and at Canjuers. No turtles are reported from Montsec. Two specimens are known from Las Hoyas. As remarked by Fabre et al. (1982), the systematics of the turtles from the Late Jurassic lithographic limestones are still somewhat uncertain : several genera have been described (Eurysternum, Idioche- lys, Solnhofia...), and they are olden included in the poorly defined family Thalassemydidae. The Las Hoyas turtles are referred to the Toxochelyi- dae by Sanz et al. (1988).

The lithographic limestone turtles are small (a few tens of centimetres in length) and usually considered as amphibious, with some some re- gression of the bony shell and limbs which have not yet been transformed into paddles. They may

SQUAMATA

They are reported from Franconia (with several genera), from Montsec (Buscalioni & Sanz 1991) and from Las Hoyas (Sanz et al. 1990). A terrestrial habitat is generally accepted.

CROCODYLIA

Metriorhynchids, which were profoundly adapted to marine life, are reported only from Francoma and from Nusplingen, with the genus Geosaurus.

Teleosaurids, which also had a mainly marine habitat, are known from Franconia and from Canjuers (Buffetaut 1980).

Atoposaurids are more widely distributed, being known from Franconia, Cerin, Montsec and Las Hoyas. There are close affinities, at the generic or even specific level, between the forms from Franconia and those from Cerin (Alligatorium, Alligatorellus, Atoposaurus, see Wellnhofer 1971), whereas the form from Montsec belongs to a different genus, Montsecosuchus {Buscalioni & Sanz

262

1990, 1991). The atoposaurids are considered as small, largely terrestrial predators.

The Crocodileimidae are represented by a single specimen of Crocodileimus robustus from Cerin. Crocodileimus was apparently a piscivorous, largely aquatic form, with, however, rather restric- ted swimming abilities (Buffetaut 1979, 1982).

Undescribed postcranlal remains from Franconia kept in the Museum ffir Naturkunde in Berlin in- dicate the existence of an additional type of croco- dilian in the Bavarian lithographic limestones. To judge from the aspect of the scutes, it may be a goniopholidid.

PTEROSAURIA

A fairly large number of pterosaur specimens is known from the Franconian localities, with several genera representatives of both the rham- phorhynchoids and the pterodactyloids (Wellnho- fer 1970, 1975). By contrast, the pterosaur record from the other localities is limited to a few isolated bones at Cerin (see Buffetaut et al. 1990) and a single skeleton at Canjuers. No pterosaurs are known the Early Cretaceous Spanish sites.

DINOSAURIA

Dinosaur skeletons are known only from Franco- nia and Canjuers, with, in each case, a single skeleton of the small theropod Compsognathus. Contrary to the opinion of Bidar et al. (1972), the Compsognathus specimen from Canjuers show no adaptations to aquatic life and must have been a terrestrial form, like its Bavarian counterpart (Ostrom 1978). An isolated theropod tooth has been reported from Las Hoyas (Frances & Sanz 1989 ; see also Buscalioni & Sanz 1991).

AVES

One of the most famous fossil vertebrates from the lithographic limestones of Franconia is doub- tless Archaeopteryx, now known by seven more or less complete skeletons and an isolated feather (see Wellnhofer 1988). No bird remains have been reported from Cerin and Canjuers.

The Spanish localities have yielded the remains of birds intermediate between Archaeopteryx and modern forms, the best skeletal material, placed in the genus Iberomesornis, coming from Las Hoyas (Sanz et al. 1988 ; Sanz 1989) ; feathers and some skeletal material are known from Montsec (Sanz et al. 1990), possibility repre- senting two genera, Ilerdopteryx and Noquerornis (Lacasa 1991).

DISCUSSION

Two main groups may be recognized among the above-mentioned lithographic limestone localities: the Late Jurassic sites of Franconia (and Nus- plingen) and Canjuers, which contain at least some marine tetrapods, and the Early Cretaceous ones of Montsec and Las Hoyas, which contain no marine tetrapods, but have yielded strictly freshwater forms (amphibians). This division is in agreement with palaeoecological reconstructions indicating connections with the open sea in Fran- conia and Provence, whereas such connections were non-existent at Las Hoyas. Although limited and sporadic contacts with the sea were once pos- tulated for Montsec (Barale et al. 1984), more recent studies suggest that no such contacts actual- ly existed and that the Montsec limestones were deposited in a shallow lacustrine environment (Mercad~ 1991 ; Martinez-Delclos et al. 1991). No truly marine tetrapods have been found at Cerin, but other evidence suggests that at least intermittent connections with the sea were possible.

There are important faunal resemblances (sometimes at the species level) between Franconia, Cerin and Canjuers. This suggests that these re- gions belonged to the same general biogeographi- cal region. Not unexpectedly, the resemblances with the more recent Spanish localities are much less obvious.

The completeness of the tetrapod record is very variable from locality to locality. The lithographic limestones of Franconia have yielded a more varied tetrapod assemblage than any other locality. They have also yielded a much larger number of fossils, although it would be difficult to give exact or even approximate figures, because the material from Franconia is scattered in a large number of collections. There is no doubt a link between the total number of fossils collected and the taxonomic diversity at each site. Fossils have been collected for more than two centuries from a large number of quarries in the Altmfihl valley, and this in itself partly explains why the Franco- nian assemblage is more diverse than those from other localities where commercial exploitation has been comparatively short-lived (about Cerin, see Bourseau et al. 1984) and where there may have been less scientific supervision.

Besides the duration of collecting activity, other factors are involved, that may account for some of the observed differences. As pointed out by various others, fossils are not particularly abundant in the lithographic limestones of Francoma, and

263

the large number "Solnhofen" fossils in many collections is mainly the result of intense quarrying activity over a long period of time. The palaeoecological excavations of the last fii~een years at Ce- rin certainly suggest that tetrapod remains are not abundant there, since only one reptile skeleton (a sphenodont) was found during that time. On the other hand, the recently discovered Las Hoyas site has already yielded a varied assemblage, containing forms which are very rare at other localities. This suggests that some lithographic limestones are clearly richer in tetrapod fossils than others.

However varied they may be, the tetrapod assemblages from the lithographic limestones of Europe do not correspond to "normal", balanced faunas. Forms which are known to have existed at the time of their deposition are conspicuously absent and, on the other hand, some of the lithographic limestone forms are unknown from other kinds of localities, or very rare there. Archaeopterix is an extreme example of the latter condition, but others can be mentioned (Crocodileimus, Compso- gnathus, etc...). Among the "missing" forms are mammals and most elements of the dinosaur fauna of the time.

Large tetrapods are very rare in the lithographic limestones of Europe. Stretosaurus was a very large animal, more than 10 metres long, but it is known from a single tooth from Francorfia. The largest tetrapod specimens from the lithographic limestones are the Steneosaurus skeletons from Franconia and Canjuers, which reach a length of some 4 metres. All large tetrapod skeletons from the lithographic limestones are those of marine forms. The lack of large dinosaur specimens is striking (the only exception being the possible carnosaur mentioned by Buscalioni & Sanz 1991, at Las Hoyas). The footprints found at Cerin in the course of the recent excavations tell a somewhat different story, however, since they show that large tetrapods visited the lagoon when its floor was out of the water or when the water was shallow. Among these animals were large turtles (Bernier et al. 1982) and, possibly, dinosaurs (Bermer et al. 1984 - but see Thulborn 1990, for another interpretation).

Fully marine tetrapods, such as ichthyosaurs or metriorhynchids, are rare in the lithographic limestones of Europe. This is to be expected in the cases of Las Hoyas and Montsec, which are interpreted as lacustrine deposits. The lack of marine forms at Cerin may suggest more difficult connections with the open sea than at Canjuers (where Steneosaurus has been found) and espe-

cially in the Franconian area, where ichthyosaurs and metriorhynchids occur.

Fully terrestrial tetrapods are represented at all sites, but nowhere can they be considered as rep- resentative of a "normal" and complete terrestrial assemblage of the Late Jurassic or Early Creta- ceous. Large forms (notably dinosaurs) are conspicuously under-represented. Other contempora- neous localities corresponding to different pa- laeoenvironments give very different pictures of tetrapod life, "dominated" by large dinosaurs, both in the Late Jurassic (fluviatile beds of the Boulonnais, for instance : see Cuny et al. 1991) and in the Early Cretaceous (Wealden of England and Belgium, non-marine Early Cretaceous of Teruel, Spain). This suggests that the terrestrial elements in the faunas from the lithographic limestones may correspond to peculiar faunal assemblages that lived in special environments, possibly close to the shores of the coastal lagoons and freshwater lakes in which the limestones were deposited.

Relatively few of the non-marine tetrapods from the lithographic limestones are known to occur at other localities corresponding to different depositional environments. Among the fbrms found elsewhere are atoposaurid crocodilians (known from the Purbeck and Wealden of England, from the Portlandian of the eastern Paris Basin, see Buffetaut, 1981, and from non-marine Early Cre- taceous deposits in Spain, see Brinkmann, 1989), the sphenodont Homoeosaurus (recently reported from "Purbeckian" deposits in Charente, France, by Buffetaut et al. 1989), and the pterosaur Cte- nochasma, also known from the Portlandmn of Haute-Marne, eastern France (Taquet 1972). This fact may reflect the already mentioned envi- ronmental bias. It may also show that our knowledge of a not negligible part of the Late Jurassic and Early Cretaceous non-marine tetrapod faunas of Europe, as reflected by non-marine localities, is incomplete, and that many of the smaller amphibians and reptiles, in particular, are under- represented. The absence of mammal remains in the lithographic limestone localities is worth mentioning.

Purely freshwater tetrapods, for which saltwater was presumably lethal, occur only in the Spanish localities, with salamanders at Las Hoyas and frogs at Montsec. The lack of amphibians in the Late Jurassic French and German localities is worth mentioning.

Flying tetrapods are very unequally represented in the lithographic limestone localities. The Fran-

264

conian localities have yielded so many specimens that they create a noticeable "Lagerst~itten effect" in the history of pterosaurs. Comparatively, the other localities are very poor in pterosaur remains. Only a few scattered isolated bones have been found at Cerin, and the reason for this is obscure (Buffetaut et al. 1990). The very small number of pterosaur remains at Canjuers may be the result of collecting over only a short period. The lack of pterosaurs in the Spanish localities, where birds occur, should also be mentioned.

The difference observed between the tetrapod assemblages from the lithographic limestones localities of Europe are not entirely due to ecological differences. It seems clear that differences in geological age are the main cause behind some differences. The Las Hoyas bird, Iberomesornis, for instance, which is more progressive than Ar- chaeopteryx, is also geologically younger, by some 25 million years.

C O N C L U S I O N S

The exceptional preservation of te trapod fossils in the Late Jurassic and Early Cretaceous lithographic l imestones of Europe is well known and has contributed greatly to the renown of these localities. It is now generally accepted that this preservation is a consequence of the peculiar environ- mental circumstances under which these sedi- ments were deposited, and various models have been put forward to account for the special sedi- mentological features of the lithographic limestones. It should not be forgotten, however, that the composition of the te trapod assemblages from the lithographic limestones is as peculiar as the preservation of the individual fossils is good. As shown by the preliminary analysis above, the tetrapod faunas from the various lithographic limestone localities in Germany, France and Spain give a picture of the te t rapod communities of the Late Jurass ic and Early Cretaceous which is different from those provided by both fully continental and fully marine localities, and is not simply the result of a mixture of both types : although they contain forms which are unknown or very poorly elsewhere, they also conspicuously lack many important elements of the faunal assemblages (both marine and continental) of the time. Whether the unusual character of these faunas is mainly the result of a taphonomic bias, or whether it does reflect the existence of peculiar communities in the areas along the shores of the lagoons and lakes in which the lithographic limestones were deposited, is still uncertain. More detailed comparative analyses of the composition of

the tetrapod assemblages from the various lithographic limestone sites in Europe may shed some new light on the interesting but still incompletely understood depositional environment of the lithographic limestones.

Acknowledgments I thank Peter Wellnhofer (Munich) and Sylvie Wenz (Paris) for providing important bibliographic references.

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Documents

Tetrapods from the Late Jurassic and Early Cretaceous lithographic limestones of Europe: A comparative review