Fabio M. Dalla Vecchia

Tethyshadros insularis, a new hadrosauroid dinosaur (Ornithischia) from the Upper Cretaceous of Italy

ABSTRACT— An articulated skeleton of a hadrosauroid dinosaur, Tethyshadros insularis n. gen., n. sp., was recovered from the Liburnian Formation (uppermost Cretaceous) of Villaggio del Pescatore in the Trieste Province of northeastern Italy. One of the most complete dinosaur fossil ever found, it shows for the first time the entire morphology of a hadrosauroid phylogenetically close to, but outside the North American and Asiatic hadrosaurids. It lived on an island developed on a carbonate platform in the Tethys Ocean and the small size of the specimens suggests that it may be an insular dwarf. The skeleton has many peculiarities including cursorial adaptations, and a mix of derived and primitive features. European hadrosauroids probably did not evolve by vicariance nor did they colonize the European Archipelago from North America, but rather came from Asia by island hopping.An articulated skeleton of a hadrosauroid dinosaur, Tethyshadros insularis n. gen., n. sp., was recovered from the Liburnian Formation (uppermost Cretaceous) of Villaggio del Pescatore in the Trieste Province of northeastern Italy. One of the most complete dinosaur fossil ever found, it shows for the first time the entire morphology of a hadrosauroid phylogenetically close to, but outside the North American and Asiatic hadrosaurids. It lived on an island developed on a carbonate platform in the Tethys Ocean and the small size of the specimens suggests that it may be an insular dwarf. The skeleton has many peculiarities including cursorial adaptations, and a mix of derived and primitive features. European hadrosauroids probably did not evolve by vicariance nor did they colonize the European Archipelago from North America, but rather came from Asia by island hopping.

A CRESTED RHAMPHORHYNCHOID PTEROSAUR FROM THE LATE TRIASSIC OF AUSTRIA

The earliest pterosaurs are represented by rare, incomplete and bearing small side cusps, posterior multicusped teeth longer specimens from the Upper Triassic (Norian) of Italy (Wild, than tall and with larger cusps, tooth size decreasing slightly 1978, 1984, 1994; Dalla Vecchia et al., 1989; Dalla Vecchia, posteriorly; tail very long without the bony sheath formed by 1994, 1995, 1998) and Greenland (Jenkins et al., 1993). The the enormously elongated preand postzygapophyses and healmost complete skeleton of a new rhamphorhynchoid pterosaur mal arches of the caudal vertebrae in other long-tailed pterois reported from the Upper Triassic of Austria. It comes from saurs (shared with Eudimorphodon). the Seefelder Schichten, which has also yielded marine fishes and terrestrial plants (Kner, 1867; Dobruskina, 1993). The enDESCRIPTION vironment of deposition of the Seefelder Schichten was a marine, anoxic and partially hypersaline basin on a very shallow The specimen is a nearly complete articulated skeleton (Fig. carbonate platform (Hopf, 1997).The main features of the spec1) preserved on a slab of black calcareous laminites. The bones imen are the presence of a sagittal cranial crest and a peculiar are preserved partly on the slab and partly on the counterslab heterodont dentition with multicusped teeth. The bony sheath, (unavailable because it was destroyed). The skull and lower consisting of very elongated caudal zygapophyses and hemal jaws broke along the sagittal plane and parts of these bones are arches, so typical of the long-tailed pterosaurs, is not present in missing. the new taxon. Cranial crests are common in Late Jurassic and The specimen is large for a Triassic pterosaur; the skull meaCretaceous pterodactyloids but have never been unambiguously sures 110 mm long and the estimated wing span is about 120 reported in rhamphorhynchoid pterosaurs. The new specimen cm. The skull is long and low, but the tip of the snout is not shows that these structures appeared very early in the history pointed (Fig. 2). The thin sagittal crest arises from the very tip of pterosaurs and must have reappeared later in pterodactyloids. of the snout and reaches its maximum height (about 20 mm) Also, Eudimorphodon has multicusped teeth but the denticulaanterior to the narial fenestra. The height of the crest decreases tion and pattern of dentition are different from the new taxon. posteriorly, as suggested by the curvature of its anterior margin The taxonomic and morphologic diversity of Triassic rhamand the distribution of the preserved fragments posteriorly. The phorhynchoids is as high as or even higher than in the succrest seems to end on the frontal directly above the orbit. It ceeding Jurassic period. becomes extremely thin at the distal (dorsal) margin, practically fading into the rocky matrix. Radial ridges are present in the anterior, highest portion. SYSTEMATIC PALEONTOLOGY The narial fenestra is elongated anteroposteriorly. The antorREPTILIA Linnaeus, 1758 bital fenestra is triangular and higher than it is long. Both are PTEROSAURIA Kaup, 1834 large openings as in Preondactylus. AUSTRIADACTYLUS CRISTATUS, gen. et sp. nov. The dentition is strongly heterodont (Fig. 3). The premaxilla (Figs. 1-3) probably bears five teeth (Fig. 3A). One or, possibly, two large teeth (10 mm high) (Fig. 3B) are present beneath the ascending Etymology--Austria, the country where the specimen was process of the maxilla. They have about 18 denticles over a found; dactylus (Greek): finger; cristatus (Latin), crested, distance of 5.5 mm along the cutting edge. The posterior maxHolotype-SMNS 56342, Staatliches Museum ftir Naturkunillary teeth (Fig. 3C) are triangular and reduced in height (4-6 de Stuttgart, Germany. mm) compared with the blade-like middle maxillary teeth. The Locality and Horizon--Abandoned mine near Ankerschlag, anterior teeth of the lower jaw were probably conical and tall. Tyrol, NW Austria; Seefelder Schichten, late Alaunian (middle The subsequent mandibular teeth are much smaller than those Norian). of the upper jaw. Seventeen multicusped teeth can be counted Diagnosis--Sagittal cranial crest extending from the tip of on the right ramus and the total number was probably 25. The the snout back to at least the middle of the orbit and deepest anterior multicusped teeth (Fig. 3D) have 4 or 5 small cusps on anterior to the naris; heterodont dentition; tall, slender conical each cutting margin. The middle mandibular teeth are slightly teeth in the premaxilla;1-2 very large, finely denticulated, smaller, leaf-shaped, and bear 6 larger cusps. The posterior blade-like teeth in the middle maxilla opposite the ascending mandibular teeth (Fig. 3E) have 4-6 cusps on each cutting process; triangular multicusped teeth with up to 12 denticles edge. along each cutting edge in the posterior part of the maxilla; Although most of the skeleton is preserved, few useful data anterior mandibular teeth similar to the premaxillary teeth, the can be discerned from the postcrania. Wing phalanx 3 is only subsequent teeth (about 25) small, leaf-shaped with 4-6 cusps slightly longer than wing phalanx 2 (103.5 mm and 101 mm on each cutting edge; anterior multicusped teeth taller than long respectively) and the ratio of their lengths (1.02) is similar to

Diversity, Relationships, and Biogeography of the Lambeosaurine Dinosaurs from the European Archipelago, with Description of the New Aralosaurin Canardia garonnensis

We provide a thorough re-evaluation of the taxonomic diversity, phylogenetic relationships, and historical biogeography of the lambeosaurine hadrosaurids from the European Archipelago. Previously published occurrences of European Lambeosaurinae are reviewed and new specimens collected from upper Maastrichtian strata of the south-central Pyrenees are described. No support is found for the recognition of European saurolophines in the available hadrosaurid materials recovered so far from this area. A new genus and species of basal lambeosaurine, Canardia garonnensis, is described on the basis of cranial and appendicular elements collected from upper Maastrichtian strata of southern France. C. garonnensis differs from all other hadrosaurids, except Aralosaurus tuberiferus, in having maxilla with prominent subrectangular rostrodorsal flange; it differs from A. tuberiferus in a few maxillary and prefrontal characters. Together with A. tuberiferus, C. garonnensis integrates the newly recognized tribe Aralosaurini. Inference of lambeosaurine interrelationships via maximum parsimony analysis indicates that the other three known European lambeosaurines are representatives of two additional subclades (tribes) of these hadrosaurids: Tsintaosaurini (Pararhabdodon isonensis) and Lambeosaurini (the Arenysaurus ardevoli-Blasisaurus canudoi clade). The tribes Aralosaurini, Tsintaosaurini, Lambeosaurini, and Parasaurolophini are formally defined and diagnosed for the first time. Three event-based quantitative methods of ancestral range reconstruction were implemented to infer the historical biogeography of European lambeosaurines: Dispersal-Vicariance Analysis, Bayesian Binary MCMC, and Dispersal-Extinction-Cladogenesis. The results of these analyses, coupled with the absence of pre-Maastrichtian lambeosaurines in the Mesozoic vertebrate fossil record of Europe, favor the hypothesis that aralosaurins and tsintaosaurins were Asian immigrants that reached the Ibero-Armorican island via dispersal events sometime during the Maastrichtian. Less conclusive is the biogeographical history of European lambeosaurins; several scenarios, occurring sometime during the Maastrichtian, are possible, from vicariance leading to the splitting of Asian or North American from European ranges to a dispersal event from North America to the European Archipelago.

First avian skeleton from the Mesozoic of northern Gondwana

Fossil birds from the Mesozoic of Gondwana are exceedingly rare (Chiappe, 1996a; Forster et al., 1998; Clarke and Chiappe, 2001). We report on a new avian specimen from the upper Cretaceous of Lebanon, which represents the only known Mesozoic bird skeleton of northern Gondwana and the first such occurrence for the African-Arabic Plate. Although recovered from marine limestones, the new fossil reveals anatomical features diagnostic of Enantiornithes, a group of Cretaceous flying birds mainly known from continental environments (Chiappe, 1995; Padian and Chiappe, 1998; Chiappe and Walker, 2002). Characteristics of its preservation and the existence of amber corpuscles scattered between its bones suggest the new Lebanese bird was transported to its marine burial from a non-marine, probably insular habitat.

THE UNUSUAL DENTITION AND FEEDING HABITS OF THE PROLACERTIFORM REPTILE LANGOBARDISAURUS (LATE TRIASSIC, NORTHERN ITALY)

Previous known specimens of the prolacertiform reptile Langobardisaurus did not allow a detailed description of the skull (Renesto, 1994a). The discovery of a new specimen from the Norian (Late Triassic) of Friuli (Northeastern Italy) allows the skull to be described for the first time, revealing a peculiar pattern of dentition for this genus, unknown in any other prolacertiform. The presence of weak, pointed anterior teeth, and bulky three-cusped cheek teeth followed posteriorly by a single, very large, grinding tooth with very small cusps, is interpreted as an adaptation to feeding on a wide variety of preys with hard exoskeletons, hard scaly covering or tough flesh. It is suggested that Langobardisaurus fed on both invertebrates and small fishes along the wide tidal flats which surrounded the anoxic basins where it was buried after death.

A vertebrate nesting site in northeastern Italy reveals unexpectedly complex behavior for late Carnian reptiles

Abstract We interpret 13 large subcircular or horseshoe-shaped depressions discovered in Late Triassic peritidal carbonate rocks of the Dogna Valley in Udine Province, northeastern Italy, to be reptile nests. These trace fossils show truncation of strata, elevated ridges of massive sediment, and sediment infill within the depression differing in shape and sedimentary structures from the host sediment. The palynological assemblage of a shaly interbed close to the nest layer indicates a Tuvalian age (late Carnian). Archosaurian footprints, produced possibly by aetosaurs, are on a surface 130 cm above the nest-bearing layer. The trackmakers are considered the most probable nest makers.

The Last Pterosaurs： First Record from the Uppermost Maastrichtian of the Tremp Syncline （Northern Spain）

: Pterosaur bones are for the first time reported from the Tremp Syncline (Spain) and the southern Pyrenees. They come from the Torrebilles-2 site (Isona, Lleida Province) in the upper Maastrichtian part of the Tremp Formation, which falls within the C29r magnetochron. Fragmentary long bones are referred to the Pterosauria based on their cortical thinness, micro-architecture, and histological features. They belong to very large to possibly giant individuals. The most complete specimen is possibly a partial right femur. The review of the Maastrichtian pterosaur record shows that only a few specimens can be referred to the C29r (which contains the K/Pg boundary). The identification of fragmentary remains could improve our knowledge about the abundance and distribution of these flying sauropsids just before the end-Cretaceous mass extinction.

A Reappraisal of the Purported Gastric Pellet with Pterosaurian Bones from the Upper Triassic of Italy.

A small accumulation of bones from the Norian (Upper Triassic) of the Seazza Brook Valley (Carnic Prealps, Northern Italy) was originally (1989) identified as a gastric pellet made of pterosaur skeletal elements. The specimen has been reported in literature as one of the very few cases of gastric ejecta containing pterosaur bones since then. However, the detailed analysis of the bones preserved in the pellet, their study by X-ray microCT, and the comparison with those of basal pterosaurs do not support a referral to the Pterosauria. Comparison with the osteology of a large sample of Middle-Late Triassic reptiles shows some affinity with the protorosaurians, mainly with Langobardisaurus pandolfii that was found in the same formation as the pellet. However, differences with this species suggest that the bones belong to a similar but distinct taxon. The interpretation as a gastric pellet is confirmed.

New information on Bobosaurus forojuliensis (Reptilia: Sauropterygia): implications for plesiosaurian evolution

Preparation of the holotype specimen of Bobosaurus forojuliensis, a large sauropterygian from the lower Carnian of northeastern Italy, revealed new morphological data relevant in establishing its phylogenetic affinities among pistosauroid taxa and its relationships with plesiosaurians. Inclusion of B. forojuliensis in two phylogenetic analyses focusing, respectively, on sauropterygians and pistosauroids agreed in placing the Italian taxon as closer to plesiosaurians than to other pistosauroids. The phylogenetic interpretation of Bobosaurus was not biased by assumptions on character weighting, is consistent with its relatively younger age compared to most pistosauroids, extends the fossil record of the plesiosaurian basal lineage back to the Carnian and supports the earliest diversification of the clade during the Late Triassic in agreement with the record of several distinct lineages of rhomaleosaurids, plesiosauroids and pliosauroids in the lowermost Jurassic. Bobosaurus shows that the evolution of the plesiosaurian body plan from the ancestral pistosauroid grade was a step-wise process, and that some of the vertebral and appendicular specialisations of Jurassic and Cretaceous plesiosaurians had already developed in the earliest Late Triassic.

THE SKULL AND LOWER JAW OF THE HOLOTYPE OF MEGALANCOSAURUS PREONENSIS (DIAPSIDA, DREPANOSAURIDAE) FROM THE UPPER TRIASSIC OF NORTHERN ITALY

The skull of the holotype of the Late Triassic diapsid reptile Megalancosaurus preonensis is described for the first time. Its study revealed affinities with archosauromorph reptiles along with striking similarities with basal pterosaurs, apart for the absence of an antorbital fenestra.