A cladistic analysis based on 39 terminal taxa and 74 characters (several multistate) using PAUP (Phylogenetic Analysis Using Parsimony) (3.1.1 for MacIntosh and 4.0b10 for Microsoft Windows) presents a new hypothesis of pterosaur inter-relationships. This study suggests that the most primitive taxon is the Anurognathidae, followed by Sordes and all remaining pterosaurs. Dendrorhynchoides is confirmed as a member of the Anurognathidae, being closely related to Batrachognathus. Preondactylus occupies a more derived position than Sordes, which questions its previous assignment as the most primitive pterosaur. The hypothesis of rhamphorhynchoid paraphyly is confirmed, with the Rhamphorhynchidae more closely related to the Pterodactyloidea than to more basal forms. The Pterodactyloidea shows a basal dichotomy: the Archaeopterodactyloidea and the Dsungaripteroidea. The Archaeopterodactyloidea is formed by Pterodactylus + Germanodactylus and a clade formed by Gallodactylidae + Ctenochasmatidae. The Nyctosauridae occupies the basal position within dsungaripteroids and is followed by the Pteranodontoidea and the Tapejaroidea. Pteranodontoids have Pteranodon at the base, followed stepwise by Istiodactylus, Ornithocheirus and the Anhangueridae. Tapejaroids are composed of the Dsungaripteridae at the base followed by the Tapejaridae and the Azhdarchidae.Major trends within pterosaur evolutionary history are: general increase in size (wing span and body); increase of wing metacarpal and pteroid; decrease of proportional length of the second and third wing phalanx relative to the first; gradual increase of rostrum (anterior to external nares); and anterior shift of the skull-mandible articulation. Cranial crests are present in most pterodactyloids, but markedly in the Ornithocheiroidea, where all taxa show some sort of crest on the skull. The loss of teeth, previously assumed to have occurred independently in several lineages, seems to be a general trend among dsungaripteroids.Several nodes recovered by this analysis are supported by very few characters, a result at least partially attributable to the limited available information from several taxa due to poor preservation and/or preparation.
Dinosauromorpha includes dinosaurs and other much less diverse dinosaur precursors of Triassic age, such as lagerpetids [1]. Joint occurrences of these taxa with dinosaurs are rare but more common during the latest part of that period (Norian-Rhaetian, 228-201 million years ago [mya]) [2, 3]. In contrast, the new lagerpetid and saurischian dinosaur described here were unearthed from one of the oldest rock units with dinosaur fossils worldwide, the Carnian (237-228 mya) Santa Maria Formation of south Brazil [4], a record only matched in age by much more fragmentary remains from Argentina [5]. This is the first time nearly complete dinosaur and non-dinosaur dinosauromorph remains are found together in the same excavation, clearly showing that these animals were contemporaries since the first stages of dinosaur evolution. The new lagerpetid preserves the first skull, scapular and forelimb elements, plus associated vertebrae, known for the group, revealing how dinosaurs acquired several of their typical anatomical traits. Furthermore, a novel phylogenetic analysis shows the new dinosaur as the most basal Sauropodomorpha. Its plesiomorphic teeth, strictly adapted to faunivory, provide crucial data to infer the feeding behavior of the first dinosaurs.
The Baurusuchidae crocodyliforms are usually interpreted as active terrestrial predators, but only some positive evidence of such habits has been described to date, mainly the relative position of external nares and orbits. Here we describe features that support this view in a complete specimen of the Baurusuchidae Stratiotosuchus maxhechti, and have executed a parsimony analysis to confirm their phylogenetic position. S. maxhechti exhibits theropodomorph features that have been previously recognized in skulls of the Baurusuchidae, as well as postcranial characteristics related to a parasagittal gait, showing that the similarities between the Baurusuchidae and theropods extend beyond the cranial morphology. These include a well‐developed supracetabular crest, a relatively medially offset femoral head and a caudally orientated calcaneal tuber. The orientations of the surfaces for muscular attachments imply that the appendicular movements of S. maxhechti were mainly anteroposterior, with abduction significantly constrained. S. maxhechti presents features that mimic some present in theropods, including a ‘fossa brevis’ on the ilium and tubercles on the ischium and femur similar to the obturator process and accessory trochanter. The relative proportions of the femur, tibia, and longer metatarsal are more similar to those of Postosuchus than to other Crocodylomorpha. In the skull, besides the theropodomorph (ziphodont) dentition concentrated in the anterior half of the rostrum, the baurusuchids are remarkable by the fusion of the nasals, which can be related to a large resistance against feeding forces acting on a high‐profile skull. The appendicular morphology of S. maxhechti strengthens the interpretation that the Baurusuchidae were active land‐dwelling predators in the Upper Cretaceous of south‐eastern Brazil, occuping ecological niches typical of small to medium‐sized theropod dinosaurs. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, S37–S56.
New specimens and an analysis of the Jehol pterosaur faunae of northeastern China show an unexpected diversity of flying reptile groups in terrestrial Cretaceous ecosystems. Here we report two new pterosaurs that are referred to European groups previously unknown in deposits of northeastern China. Feilongus youngi, from the Yixian Formation, is closely related to the Gallodactylidae and is distinguished by the presence of two independent sagittal crests and a protruding upper jaw. Nurhachius ignaciobritoi, from the Jiufotang Formation, has teeth formed by labiolingually compressed triangular crowns, only previously reported in Istiodactylus latidens from England. With these new discoveries, the Jehol pterosaurs show a wide range of groups including both primitive and derived forms that are not matched by any other deposit in the world. The discoveries also document the turnover of pterosaur faunae, with the primitive Anurognathidae and early archaeopterodactyloids being replaced by derived pterodactyloids. Furthermore, these deposits offer an opportunity to examine the interaction and competition between birds and pterosaurs--it indicates that the avian fauna during the Lower Cretaceous (and possibly most of the Mesozoic) dominated terrestrial, inland regions, whereas pterosaurs were more abundant in coastal areas.
A pterosaur bone bed with at least 47 individuals (wing spans: 0.65–2.35 m) of a new species is reported from southern Brazil from an interdunal lake deposit of a Cretaceous desert, shedding new light on several biological aspects of those flying reptiles. The material represents a new pterosaur, Caiuajara dobruskii gen. et sp. nov., that is the southermost occurrence of the edentulous clade Tapejaridae (Tapejarinae, Pterodactyloidea) recovered so far. Caiuajara dobruskii differs from all other members of this clade in several cranial features, including the presence of a ventral sagittal bony expansion projected inside the nasoantorbital fenestra, which is formed by the premaxillae; and features of the lower jaw, like a marked rounded depression in the occlusal concavity of the dentary. Ontogenetic variation of Caiuajara dobruskii is mainly reflected in the size and inclination of the premaxillary crest, changing from small and inclined (∼115°) in juveniles to large and steep (∼90°) in adults. No particular ontogenetic features are observed in postcranial elements. The available information suggests that this species was gregarious, living in colonies, and most likely precocial, being able to fly at a very young age, which might have been a general trend for at least derived pterosaurs.
A new long-tailed pterosaur, Wukongopterus lii gen. et sp. nov, is described based on an almost complete skeleton (IVPP V15113) representing an individual with an estimated wing span of 730 mm. The specimen was discovered in strata that possibly represent the Daohugou Bed (or Daohugou Formation) at Linglongta, Jianchang, Liaoning Province, China. Wukongopterus lii is a non-pterodactyloid pterosaur diagnosed by the first two pairs of premaxillary teeth protruding beyond the dentary, elongated cervical vertebrae (convergent with Pterodactyloidea), and a strongly curved second pedal phalanx of the fifth toe. The specimen further has a broken tibia that indicates an injury occurred while the individual was still alive. Taphonomic aspects provide indirect evidence of an uropatagium, supporting the general hypothesis that at least all non-pterodactyloid pterosaurs show a membrane between the hind limbs. A phylogenetic analysis including most non-pterodactyloid pterosaurs shows that Wukongopterus lii gen. et sp. nov. lies outside the Novialoidea, being cladistically more primitive than the Rhamphorhynchidae and Campylognathoides. This analysis differs from previous studies and indicates that more work is needed before a stable picture of non-pterodactyloid pterosaur relationships is achieved.
Considered one of the best known flying reptiles, Pteranodon has been subject to several reviews in the last century. Found exclusively in the Upper Cretaceous Niobrara Formation and Pierre Shale Group 11 species have been attributed to this genus (excluding the ones presently regarded as representing Nyctosaurus). While reviewers agree that this number is inflated, there is disagreement on how many species can be identified. The last review recognized only two species (Pteranodon longiceps and Pteranodon sternbergi) both being sexually dimorphic. Based on several cranial features, some specimens previously referred to the genus Pteranodon are re-evaluated leading to the recognition of the following species, two of which new that are described here: Pteranodon longiceps, Geosternbergia sternbergi, Geosternbergia maiseyi sp. nov., and Dawndraco kanzai gen. et sp. nov. They differ mainly by features such as the direction and extension of the frontal crest, the angle and extension of the posterior process of the premaxillae, the shape and extension of the lower temporal fenestra and the length and proportion of the rostrum. The procedures to recognize a pterosaur species are also discussed here, and must take into account primarily morphology, in conjunction with stratigraphic and geographic data. Although well aware that changes in morphology not always reflect taxonomy, the lack of stratigraphic data and the limited number of specimens that can be confidently assigned to one species hampers our understanding on the morphological variations as a function of ontogeny, individual variation and sexual dimorphism. Although the present study has not eliminated the possibility to recognize such differences, caution is needed before models are generalized for pterosaurs.
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