Deux nouveaux sites à traces de pas dinosauroïdes dans la Formation Dolomitique de l’Hettangien de Lozère,Languedoc-Roussillon,France

Recent ichnological studies revealed two new localities showing dinosaur footprints in the Hettangian Dolomitic Formation from the Causses Basin, southern France. The traces are reported from the northern part of the basin, in Lozère, an area where Lower Jurassic dinosaur ichnites were poorly documented. The surfaces bearing footprints are characterized by a large number of traces. The ichnites are ascribed to Grallator (Hitchcock, 1858), Dilophosauripus (Welles, 1971) and Eubrontes (Hitchcock, 1845) whose trackmakers were theropods. Although numerous tracksites were previously described in the southern part of the Causses Basin, these three ichnotaxa are for the first time observed in association into the Dolomitic Formation.     

Large Carnivore Footprints from the Late Pleistocene of Argentina

The aim of the present contribution is to describe large felipedid footprints from a new ichnological site from the Late Pleistocene of Buenos Aires Province, Argentina. The prints are referred as the new ichnospecies Felipeda miramarensis nov. ichnosp. Based on size and morphology, this new ichnotaxon may have been produced by the large machairodontine felid Smilodon populator. Track analysis indicates that the producer of the tracks had fully retractile claws, a plantigrade feet, and lacked strong supination capabilities on pes and manus. The size and depth differences between manus and pes prints indicate that the producer had notably robust anterior limbs. If correctly assigned, the new ichnospecies reinforces the idea that Smilodon was an ambush predatory mammal.     

On the Procompsognathus postcranium (Late Triassic, Germany)

A review of the historical background of the material housed in the Staatliches Museum für Naturkunde (Stuttgart) and ascribed to Procompsognathus triassicus (Upper Triassic, Germany) is provided. The systematic position of the postcranial remains is discussed. The combined results of cladistic analyses suggest that the type material, an incomplete postcranial skeleton in two pieces (SMNS 12591), is from a theropod close to Segisaurus and Coelophysis. An isolated manus (SMNS 12352a) is definitely not theropodan, but could be from any small basal archosaur. The remarkable diversity of the carnivorous guild that dwelled in southern Germany before the end-Triassic events is underlined.     

Chirothere Footprint Sites from the Otter Sandstone Formation (Middle Triassic,late Anisian) of Devon,United Kingdom

Three chirothere footprint sites are documented from the Middle Triassic (Anisian) fluvial Otter Sandstone Formation of Sidmouth, Devon, UK. One site, on the foreshore below Peak Hill, has revealed numerous tracks, including a probable manus imprint, which are recorded here. Another site, also beneath Peak Hill, has produced a single probable manus track from a higher horizon. The third site, on the foreshore below Salcombe Hill Cliff, has yielded a small number of pes tracks similar to those from Peak Hill. The footprints at all sites are interpreted as belonging to a large Chirotherium ichnospecies, perhaps C. barthii Kaup. Unusually, the tracks are preserved in mudstone, often in convex epirelief, and are inferred to have been generated subaqueously. The apparent restriction of the footprints to the higher part of the Otter Sandstone Formation supports other evidence, such as a greater abundance of vertebrate skeletal material, subhorizontal rather than vertical rhizocretions, and a greater frequency of lacustrine facies, for a less arid climate and higher water table than lower in the sequence.     

Late Triassic Aetosaurs as the Trackmaker of the Tetrapod Footprint Ichnotaxon Brachychirotherium

Brachychirotherium is the common ichnogenus of Late Triassic chirothere footprints well known from western Europe, North America, Argentina and South Africa. Although it has long been agreed by most workers that the trackmaker of Brachychirotherium was a derived crurotarsan archosaur, the trackmaker has been identified as either a rauisuchian or an aetosaur, and some workers attribute it to a primitive crocodylomorph (sphenosuchian). New knowledge of the osteology of the manus and pes of a large aetosaur, Typothorax coccinarum, indicates a close correspondence between the manus and pes structure of aetosaurs and the morphology of Brachychirotherium. Furthermore, functional analysis of complete skeletons indicates aetosaurs plausibly placed their feet in the narrow gauge, nearly the overstepped walk characteristic of Brachychirotherium. Brachychirotherium and aetosaurs have matched distributions, that is, they were Pangea-wide during the Late Triassic. The manus and pes morphology of rauisuchians and early crocodylomorphs (sphenosuchians) deviate from Brachychirotherium footprint morphology in key features, thus excluding their identification as trackmakers. Aetosaurs made Brachychirotherium footprints.     

New Permian tetrapod footprints and macroflora from Turkey (Çakraz Formation, northwestern Anatolia): Biostratigraphic and palaeoenvironmental implications

New tetrapod footprints belonging to the ichnogenus Hyloidichnus have been discovered in Turkey for the first time, in the lower part of the Çakraz Formation (Northwestern Anatolia) and together with macrofloral imprints of Annularia and Stigmaria. These discoveries confirm the Permian age of the fossiliferous red beds in which the coniferophyte Walchia was previously recorded. Based on the stratigraphic range of Annularia, Stigmaria and Hyloidichnus known elsewhere, a Cisuralian age is proposed for these beds. These new ichno- and macrofloral remains, together with the sedimentological data (mudcracks, rain drops) suggest the presence of captorhinid reptiles living in a palustrine floodplain environment, and under a warm, seasonal climate alternating between humid and relatively long dry seasons. These climatic conditions may have permitted the migration of these captorhinids through Laurasia during the Permian.     

A Revised Systematic Ichnotaxonomy and Review of the Vertebrate Footprint Ichnofamily Chirotheriidae from the British Triassic

Since 1838, many discoveries of the tetrapod footprint taxon Chirotherium and closely similar forms have been reported from the British Middle Triassic. These have been assigned to dozens of ichnotaxa, so that the identification and interpretation of members of the ichnofamily Chirotheriidae have become confused.

Part 1 of this article deals with ichnotaxonomy. Revision of all the available material shows that there are at least three valid ichnospecies of Chirotherium, one of Synaptichnium, and one of Isochirotherium. The presence of a fourth related ichnogenus, Brachychirotherium, cannot yet be confirmed in Britain. Emended diagnoses are proposed for all four ichnogenera; they may be readily distinguished by the shape of the hand (manus) and foot (pes) impression and especially by the phalangeal counts. The natural cast of a probable tail impression, associated with Chirotheriidae footprints from the British Triassic is figured for the first time, as is a rare detailed impression of skin and the tarsal region of an Isochirotherium lomasi pes cast.

Part 2 provides a critical history of the earliest discoveries of each of these ichnogenera in the British Triassic. It is subdivided into divisions, depending on the quality of data and existence of specimens, the localities in Britain which have yielded these ichnogenera; a detailed history and discussion is provided of the earliest discoveries made at each locality. The stratigraphic distribution and paleological significance of the ichnogenera/species are assessed.     

Were some dinosaurs gregarious?

A careful survey of the dinosaur footprints and trackways (identified as Eubrontes, Anchisauripus and Grallator) at the Mt. Tom site north of Holyoke, Massachusetts, and plots of their orientations reveal that an improbable percentage (70%) of the tracks are oriented in a nearparallel course. All but one of these coincident ancient traverses were probably made by the same kind of Triassic dinosaur (the footprints of which are referred to as Eubrontes). Comparable trackway orientation patterns have been reported by Albritton (1942) near Comanche, Texas for an Early Cretaceous dinosaur (iguanodobntid?) and Bird (1941, 1944) cited paralled sauporod trackways of Early Cretaceous age near the Paluxy River in Texas. Probability dictates that these sub-parallel traverses were not independent events and the presence of other deviating trackways at all three sites indicates that the trackmakers probably were not confined in their passage by physical barriers. Furthermore, the coincidental occurrence of such natural barriers at all of the sites mentioned here seems highly improbable. the combinedevidence of the Massachusetts site and the two Texas localities, together with the apparently widespread occurrence of dinosaur trackway lineation, strongly indicates gregarious habits for several different kinds of dinosaurs.     

History of Ichnology: The Misconceived Footprints of Rhynchosaurs

In 1842 Richard Owen described a Triassic reptile from Grinshill, Shropshire, which he named Rhynchosaurus articeps. He suggested that footprints found in the same beds were those of this fossil. However, the footprints were characterised by a backward-pointing toe and so were of the type now known as Rotodactylus. Huxley (1877), Woodward (1907), and Benton (1990) have subsequently shown that the five digits of Rhynchosaurus point forward and so could not have left these footprints. In 1896 Beasley classified the Triassic footprints found in Cheshire, his type D prints being those earlier assigned to rhynchosaurs. His D1 prints were named Rhynchosauroides articeps by Maidwell (1911). However, these D1 prints, which come from a lower horizon in the Anisian, are consistently too small to match Owen's fossil. Beasley's D3 form, now named Synaptichnium pseudosuchoides Nopcsa, is more likely to represent the footprints of Rhynchosaurus articeps, although further research and study of more complete trackways will be necessary to clarify whether these are the footprints of Archosauromorphs, such as rhynchosaurs or possibly those of Archosauriformes, for example, erythrosuchids or proterosuchids. Maidwell's Rhynchosauroides rectipes and Rhynchosauroides membranipes, originally believed to be distinct ichnospecies, are more likely to be synonyms, their apparent differences reflecting variations in the substrate traversed.     

Paleocene Thalassinidea colonization in deep-sea environment and the coprolite Palaxius osaensis n. ichnosp. in Southern Costa Rica

Palaxius osaensis n. ichnosp., a new ichnospecies of crustacean coprolite is described. The coprolite is preserved in a 200-m thick Paleocene sequence in Southern Costa Rica that is largely dominated by pillow basalts. The studied sample is part of a seamount formed in the Pacific Ocean that was accreted to the Central American isthmus during the Eocene. The absence of lava vesicles, shallow-water deposits, and detrital sediments in the section suggest that the coprolites were deposited in a deep environment during the first stage of the development of the seamount. This represents one of the deepest occurrences of Thalassinidea coprolites reported in the literature and indicates that the producers of the coprolites, presumably some shrimps, developed the aptitude to colonize abyssal environments at least since Early Tertiary. The crustacean coprolites were encountered at a site which apparently lacked a food supply, although hydrothermal processes are believed to have provided the opportunity for a chemotrophic community to develop on the deepest part of the seamount. P. osaensis n. ichnosp. is also found in Colombia in late Cretaceous shallow-water sediments that notably contain Palaxius caucaensis coprolites (Micropaleontology 41 (1995) 85-88). Occurrences of P. osaensis n. ichnosp. deposited at both shallow and deep levels may possibly be related to an aptitude of some thalassinid organisms to have developed in various biotopes during the late Cretaceous-Paleocene.     

First Early Jurassic Ornithischian and theropod footprint assemblage and a new ichnotaxon Shenmuichnus wangi ichnosp. nov. from Yunnan Province,southwestern China

The new ichnospecies, Shenmuichnus wangi ichnosp. nov., is the first evidence for the presence of large ornithischians in the Early Jurassic of Yunnan Province, whereas the known skeletal record documents small species only. Until now Shenmuichnus was known from a single locality in Shaanxi Province by the ichnospecies Shenmuichnus youngteilhardorum. Compared with the latter, Shenmuichnus wangi is larger and shows a different trackway configuration, particularly in the relative position of manus and pes imprints. Palecologically, the occurrence of Shenmuichnus wangi in a red bed facies indicates the preference of distinctive environments of trackmakers of both ichnospecies, questioning former hypotheses of exclusivity of ornithischians in more humid climates. By abundance both skeletons and footprints of ornithischians suggest their role as a minor component in Early Jurassic saurischian dominated dinosaur faunas in this region.     

Reinvestigation of conchostracans (Crustacea: Branchiopoda) from the Permian–Triassic transition in Southwest China

Sedimentary deposits of the Permian–Triassic transition are well-exposed in numerous outcrops of South China. Depending on the palaeogeographic positions of the sections, their lithofacies vary from fully marine, shallow marine, lagoonal, lacustrine, and fluvial to alluvial. In the present study, conchostracans (Crustacea: Branchiopoda) were newly collected from the continental deposits of the Kayitou and Jialingjiang formations around the Kangdian Highland elevated by the Emeishan Basalt in the southern Sichuan, western Guizhou, and northeastern Yunnan provinces. The conchostracan fauna of the Kayitou Formation is composed of Pseudestheria chatangensis, Euestheria fuyuanensis, and Euestheria sp. aff. E. gutta. These species occur in varying lithofacies types of different sections. In particular, the late Permian Pseudestheria chatangensis occurs in grey siltstones interbedded with pebbly sandstones, which are interpreted as lacustrine deposits. Euestheria fuyuanensis and Euestheria sp. aff. E. gutta were obtained from yellowish to greenish–grey clay- and siltstones, interpreted as coastal plain deposits. In comparison to other regions, occurrences of Euestheria gutta are indicative of an early Induan to Olenekian (Early Triassic) age. The fossil record of Euestheria fuyuanensis is so far restricted to a few occurrences in the Kayitou Formation of Southwest China and, therefore, using this species for long-distance biostratigraphic correlation requires further investigation. The distribution of late Permian pseudestheriid and Early Triassic euestheriid species in the respective sections possibly depends on the local lithofacies and, therefore, a diachronous age of the Kayitou Formation within the study area can be assumed. Additionally, Magniestheria sp. aff. M. mangaliensis and Magniestheria sp. aff. M. subcircularis were recorded in the Jialingjiang Formation, which represents a lithostratigraphic unit considered as late Early Triassic (Olenekian). Further investigations of both taxonomy and the real stratigraphic ranges of the conchostracan species as well as cross-correlations with other age data are recommended, in order to better constrain the position of the Permian–Triassic boundary and the specific timing of the terrestrial end-Permian mass extinction in continental deposits of Southwest China.     

Refinements in the Upper Permian to Lower Jurassic stratigraphy of Karakorum, Pakistan

New sampling on critical intervals of the uppermost Permian and Triassic successions of the Northern Karakorum Terrain in the Karakorum Range (Pakistan) has refined the stratigraphy. Two types of successions may be distinguished in the Karakorum Range: a carbonate platform succession, spanning the whole interval from Upper Permian to Upper Triassic, possibly with several gaps; and a basinal succession, deposited from the Middle Permian to Early Carnian (Late Triassic), when the carbonate platform prograded into the basin. With the approaching and later docking of the Karakorum Block against the Asian margin closing the Paleo-Tethys, a portion of Karakorum emerged while another part subsided as a fore-deep, receiving clastics from the emerging Cimmerian Range. Molassic sediments filled the basin, whilst shallow-water carbonates transgressed over the emerged carbonate platform sometime between the latest Triassic and the Pliensbachian (Early Jurassic), with Cimmerian deformation occurring to the north. The age control is provided by conodonts, with assemblages of late Wuchiapingian, Changhsingian, Induan (Griesbachian and Dienerian), late Olenekian, early Anisian, late Ladinian, and early Carnian ages, respectively. Some information on the section around the P/T boundary is provided by palynology and isotopic C¹³ values. The dating of the Norian/Rhaetian platform is provided by foraminifers.     

Diaceratherium massiliae nov. sp. des argiles oligocènes de Saint-André et Saint-Henri à Marseille et de Les Milles près d’Aix-en-Provence (SE de la France), premier grand Rhinocerotidae brachypode européen

A revision of all previously collected mammalian fossils from the two Late Oligocene sites of Saint-André and Saint-Henri in Marseille (both from the MP 26 reference-level) allows us to identify three Rhinocerotoidea species: Protaceratherium albigense, Ronzotherium romani, Diaceratherium massiliae nov. sp., and maybe a fourth one, Eggysodon cf. gaudryi. Only the first two were previously known there. D. massiliae nov. sp. is found together with R. romani; it is the first case of sympatry ever known between the two genera. D. massiliae nov. sp. is then the most ancient Diaceratherium in Europe, where the genus was previously unknown before the MP 29 reference-level. It is a very large species whose limb bones proportions foreshadow these of the later species of the genus, especially D. lemanense from the Late Oligocene-Early Miocene. This suggests a possible phylogenetic link between D. massiliae nov. sp. and D. lemanense, and the coexistence of at least two different but partially contemporaneous lineages among the European Diaceratherium. In Les Milles near Aix-en-Provence, also from the MP 26 reference-level, the three species P. albigense, R. romani and D. massiliae nov. sp. were also found.     

Juvenile ornithopod (Dinosauria: Rhabdodontidae) remains from the Upper Cretaceous (Lower Campanian, Gosau Group) of Muthmannsdorf (Lower Austria)

The fragmentary remains of a juvenile rhabdodontid ornithopod from the Coal-bearing Complex of the Gosau Group (Lower Campanian, Grünbach syncline) at Muthmannsdorf near Wiener Neustadt, Lower Austria are revised. The material, probably belonging to a single individual, includes a right dentary (lectotype of Iguanodon suessi Bunzel, 1871, designated herein), teeth, a fragmentary parietal, fragments of scapula, ?radius, femur, tibia, two vertebrae (lost) and a manual ungual.The lectotype dentary does not provide clear autapomorphies or sufficient diagnostic features to determine its position within the Rhabdodontidae at generic level. By this “Iguanodon suessi” Bunzel, 1871 and the genus “Mochlodon” Seeley, 1881, to which it was latter referred as type species, cannot be characterized sufficiently by differential diagnosis and these are best considered nomina dubia. Based upon combined character comparisons (mainly postcranial features) the Muthmannsdorf ornithopod is referred herein to Zalmoxes Weishampel, Jianu, Csiki and Norman, 2003, a genus so far known from the late Maastrichtian of Romania. It probably but not evidently represents a yet unnamed species, most closely related to Zalmoxes shqiperorum Weishampel, Jianu, Csiki and Norman, 2003. At the present state of knowledge the Austrian material is not further diagnostic at the species level and kept in open nomenclature as Zalmoxes sp.     

Uniquely derived upper molar morphology of Eocene Amphipithecidae (Primates: Anthropoidea): Homology and phylogeny

The extinct Southeast Asian primate family Amphipithecidae is regularly cited in discussions of anthropoid origins, but its phylogenetic position remains controversial. In part, the lack of consensus regarding amphipithecid relationships can be attributed to uncertainty regarding the homology of upper molar structures in this group. Here, we describe a virtually pristine upper molar of Pondaungia cotteri from the late middle Eocene Pondaung Formation of Myanmar, which is the first example of a relatively unworn and well-preserved amphipithecid upper molar ever recovered. The distolingual upper molar cusp in this new specimen of Pondaungia appears to be a lingually displaced and enlarged metaconule, rather than a hypocone or pseudohypocone as previous workers have thought. Reassessment of the upper molar morphology of other amphipithecids and putative amphipithecids reveals a very similar pattern in Siamopithecus, Myanmarpithecus and Ganlea, all of which are interpreted as having upper molars showing many of the same derived features apparent in Pondaungia. In contrast, the upper molar morphology of Bugtipithecus diverges radically from that of undoubted amphipithecids, and the latter taxon is excluded from Amphipithecidae on this basis. Phylogenetic analyses of several character–taxon matrices culled from the recent literature and updated to reflect the new information on amphipithecid upper molar morphology yield similar results. Consensus tree topologies derived from these analyses support amphipithecid monophyly and stable relationships within Amphipithecidae. Amphipithecids appear to be stem members of the anthropoid clade.     

Biometric and morphometric approaches on Lower Hettangian dinosaur footprints from the Rodez Strait (Aveyron,France)

Along the southern crystalline border of the Rouergue, the detrital Sandstones-variegated Mudstones Formation and the Dolomitic Formation yields numerous dinosaur footprints from the Earliest Hettangian. Among the 25 sites distributed along the 40 km-long transect between Marcillac-Vallon and Saint-Geniez-d’Olt, two of them, Puech de Castres and Le Bouyssou, have yielded abundant ichnites in various well-defined stratigraphic units. The combined analysis of these footprints with biometric and morphometric methods (Fourier analysis) warrants their identification to Grallator, Eubrontes and Dilophosauripus. The dual methodological approach also reveals two new morphotypes previously unknown in the Causses Basin. Moreover, the footprints assigned to Grallator and Eubrontes tend to be morphologically similar, suggesting that they might correspond to a same taxon of Coelophysoidea theropod at different growth stages. This dinosaur fauna was living in a vast floodplain, in which fluvial channels carried southward the detrital material coming from the Rouergue substratum. The Dolomitic Formation was deposited in the context of a northward marine transgression during which shallow and aerial environments prevailed in the area.     

Ichnotaxonomic Review of Large Ornithopod Dinosaur Tracks: Temporal and Geographic Implications

BackgroundLarge ornithopod tracks are known from the Upper Jurassic to the uppermost Cretaceous rocks of all continents but Antarctica. They include the tracks historically called Iguanodon footprints, iguanodontid footprints, hadrosaur/hadrosaurid footprints, and other large ornithopod tracks that have been used to define ichnotaxa. More than 40 ichnospecies based on large ornithopod tracks have been defined, but the validity of many of them is questionable.Conclusions/SignificanceThe monospecific ichnogenus Iguanodontipus (I. burreyi) is characterized by a small, rounded heel and elongate, narrow digit impressions. Its distribution is limited to the Berriasian-Valanginian of Europe. Caririchnium consists of four ichnospecies (C. magnificum [type ichnospecies], C. kortmeyeri, C. billsarjeanti and C. lotus) with a large, rounded heel and short, wide digit impressions. This ichnogenus ranges from the Berriasian-Hauterivian to the Aptian-Albian of South America, North America, Asia and Europe. Finally, Hadrosauropodus (three ichnospecies: H. langstoni [type ichnospecies], H. leonardii and H. kyoungsookimi) shows a large, bilobed heel and short, wide digit impressions. It is known from the Aptian-Albian to the Maastrichtian of North America, Asia and Europe. The ichnofamily Iguanodontipodidae includes large iguanodontian tracks characterized mainly by mesaxonic, tridactyl and subsymmetrical pes tracks that are as wide as (or wider than) long and have one pad impression in each digit and one in the heel. Its distribution is confidently limited to the Cretaceous of Europe, Asia, North America and South America.