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Concept: Saurischia


Relationships between non-avian theropod dinosaurs and extant and fossil birds are a major focus of current paleobiological research. Despite extensive phylogenetic and morphological support, behavioural evidence is mostly ambiguous and does not usually fossilize. Thus, inferences that dinosaurs, especially theropods displayed behaviour analogous to modern birds are intriguing but speculative. Here we present extensive and geographically widespread physical evidence of substrate scraping behavior by large theropods considered as compelling evidence of “display arenas” or leks, and consistent with “nest scrape display” behaviour among many extant ground-nesting birds. Large scrapes, up to 2 m in diameter, occur abundantly at several Cretaceous sites in Colorado. They constitute a previously unknown category of large dinosaurian trace fossil, inferred to fill gaps in our understanding of early phases in the breeding cycle of theropods. The trace makers were probably lekking species that were seasonally active at large display arena sites. Such scrapes indicate stereotypical avian behaviour hitherto unknown among Cretaceous theropods, and most likely associated with terrirorial activity in the breeding season. The scrapes most probably occur near nesting colonies, as yet unknown or no longer preserved in the immediate study areas. Thus, they provide clues to paleoenvironments where such nesting sites occurred.

Concepts: Bird, Reptile, Fossil, Dinosaur, Saurischia, Theropoda, Maniraptora, Avialae


Late Cretaceous terrestrial strata of the Neuquén Basin, northern Patagonia, Argentina have yielded a rich fauna of dinosaurs and other vertebrates. The diversity of saurischian dinosaurs is particularly high, especially in the late Cenomanian-early Turonian Huincul Formation, which has yielded specimens of rebacchisaurid and titanosaurian sauropods, and abelisaurid and carcharodontosaurid theropods. Continued sampling is adding to the known vertebrate diversity of this unit.

Concepts: Bird, Cretaceous, Dinosaur, Saurischia, Theropoda, Ceratosauria, Late Cretaceous, Carnosauria


Oviraptorosaurs are a bizarre group of bird-like theropod dinosaurs, the derived forms of which have shortened, toothless skulls, and which diverged from close relatives by developing peculiar feeding adaptations. Although once among the most mysterious of dinosaurs, oviraptorosaurs are becoming better understood with the discovery of many new fossils in Asia and North America. The Ganzhou area of southern China is emerging as a hotspot of oviraptorosaur discoveries, as over the past half decade five new monotypic genera have been found in the latest Cretaceous (Maastrichtian) deposits of this region. We here report a sixth diagnostic oviraptorosaur from Ganzhou, Tongtianlong limosus gen. et sp. nov., represented by a remarkably well-preserved specimen in an unusual splayed-limb and raised-head posture. Tongtianlong is a derived oviraptorid oviraptorosaur, differentiated from other species by its unique dome-like skull roof, highly convex premaxilla, and other features of the skull. The large number of oviraptorosaurs from Ganzhou, which often differ in cranial morphologies related to feeding, document an evolutionary radiation of these dinosaurs during the very latest Cretaceous of Asia, which helped establish one of the last diverse dinosaur faunas before the end-Cretaceous extinction.

Concepts: Skull, Cretaceous, Dinosaur, Saurischia, Theropoda, Maniraptora, Oviraptorosauria, Oviraptoridae


Titanosauria is an exceptionally diverse, globally-distributed clade of sauropod dinosaurs that includes the largest known land animals. Knowledge of titanosaurian pedal structure is critical to understanding the stance and locomotion of these enormous herbivores and, by extension, gigantic terrestrial vertebrates as a whole. However, completely preserved pedes are extremely rare among Titanosauria, especially as regards the truly giant members of the group. Here we describe Notocolossus gonzalezparejasi gen. et sp. nov. from the Upper Cretaceous of Mendoza Province, Argentina. With a powerfully-constructed humerus 1.76 m in length, Notocolossus is one of the largest known dinosaurs. Furthermore, the complete pes of the new taxon exhibits a strikingly compact, homogeneous metatarsus-seemingly adapted for bearing extraordinary weight-and truncated unguals, morphologies that are otherwise unknown in Sauropoda. The pes underwent a near-progressive reduction in the number of phalanges along the line to derived titanosaurs, eventually resulting in the reduced hind foot of these sauropods.

Concepts: Sauropoda, Dinosaur, Sauropodomorpha, Diplodocus, Saurischia, Titanosaur, Bruhathkayosaurus, Macronaria


Medullary bone (MB), an estrogen-dependent reproductive tissue present in extant gravid birds, is texturally, histologically and compositionally distinct from other bone types. Phylogenetic proximity led to the proposal that MB would be present in non-avian dinosaurs, and recent studies have used microscopic, morphological, and regional homologies to identify this reproductive tissue in both theropod and ornithischian dinosaurs. Here, we capitalize on the unique chemical and histological fingerprint of MB in birds to characterize, at the molecular level, MB in the non-avian theropod Tyrannosaurus rex (MOR 1125), and show that the retention of original molecular components in fossils allows deeper physiological and evolutionary questions to be addressed.

Concepts: Molecular biology, Bird, Paleontology, Dinosaur, Saurischia, Theropoda, Cretaceous–Tertiary extinction event, Tyrannosaurus


For 130 years, dinosaurs have been divided into two distinct clades-Ornithischia and Saurischia. Here we present a hypothesis for the phylogenetic relationships of the major dinosaurian groups that challenges the current consensus concerning early dinosaur evolution and highlights problematic aspects of current cladistic definitions. Our study has found a sister-group relationship between Ornithischia and Theropoda (united in the new clade Ornithoscelida), with Sauropodomorpha and Herrerasauridae (as the redefined Saurischia) forming its monophyletic outgroup. This new tree topology requires redefinition and rediagnosis of Dinosauria and the subsidiary dinosaurian clades. In addition, it forces re-evaluations of early dinosaur cladogenesis and character evolution, suggests that hypercarnivory was acquired independently in herrerasaurids and theropods, and offers an explanation for many of the anatomical features previously regarded as notable convergences between theropods and early ornithischians.

Concepts: Bird, Phylogenetics, Dinosaur, Sauropodomorpha, Saurischia, Theropoda, Coelurosauria, Ornithischia


Theropod dinosaurs were the dominant predators in most Mesozoic era terrestrial ecosystems. Early theropod evolution is currently interpreted as the diversification of various carnivorous and cursorial taxa, whereas the acquisition of herbivorism, together with the secondary loss of cursorial adaptations, occurred much later among advanced coelurosaurian theropods. A new, bizarre herbivorous basal tetanuran from the Upper Jurassic of Chile challenges this conception. The new dinosaur was discovered at Aysén, a fossil locality in the Upper Jurassic Toqui Formation of southern Chile (General Carrera Lake). The site yielded abundant and exquisitely preserved three-dimensional skeletons of small archosaurs. Several articulated individuals of Chilesaurus at different ontogenetic stages have been collected, as well as less abundant basal crocodyliforms, and fragmentary remains of sauropod dinosaurs (diplodocids and titanosaurians).

Concepts: Bird, Reptile, Cretaceous, Dinosaur, Saurischia, Archaeopteryx, Theropoda, Jurassic


A monodominant bonebed of Avimimus from the Nemegt Formation of Mongolia is the first oviraptorosaur bonebed described and the only recorded maniraptoran bonebed from the Late Cretaceous. Cranial elements recovered from the bonebed provide insights on the anatomy of the facial region, which was formerly unknown in Avimimus. Both adult and subadult material was recovered from the bonebed, but small juveniles are underrepresented. The taphonomic and sedimentological evidence suggests that the Avimimus bonebed represents a perimortem gregarious assemblage. The near absence of juveniles in the bonebed may be evidence of a transient age-segregated herd or ‘flock’, but the behaviour responsible for this assemblage is unclear. Regardless, the Avimimus bonebed is the first evidence of gregarious behaviour in oviraptorosaurs, and highlights a potential trend of increasing gregariousness in dinosaurs towards the end of the Mesozoic.

Concepts: Bird, Reptile, Cretaceous, Dinosaur, Saurischia, Theropoda, Maniraptora, Oviraptorosauria


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.

Concepts: Fossil, Dinosaur, Sauropodomorpha, Saurischia, Triassic, Carnian, Archosaur, Norian


Maniraptora includes birds and their closest relatives among theropod dinosaurs. During the Cretaceous period, several maniraptoran lineages diverged from the ancestral coelurosaurian bauplan and evolved novel ecomorphologies, including active flight, gigantism, cursoriality and herbivory. Propagation X-ray phase-contrast synchrotron microtomography of a well-preserved maniraptoran from Mongolia, still partially embedded in the rock matrix, revealed a mosaic of features, most of them absent among non-avian maniraptorans but shared by reptilian and avian groups with aquatic or semiaquatic ecologies. This new theropod, Halszkaraptor escuilliei gen. et sp. nov., is related to other enigmatic Late Cretaceous maniraptorans from Mongolia in a novel clade at the root of Dromaeosauridae. This lineage adds an amphibious ecomorphology to those evolved by maniraptorans: it acquired a predatory mode that relied mainly on neck hyperelongation for food procurement, it coupled the obligatory bipedalism of theropods with forelimb proportions that may support a swimming function, and it developed postural adaptations convergent with short-tailed birds.

Concepts: Bird, Dinosaur, Saurischia, Archaeopteryx, Theropoda, Coelurosauria, Dromaeosauridae, Maniraptora