A new avialan theropod from an emerging Jurassic terrestrial fauna.

Birds are descended from non-avialan theropod dinosaurs of the Late Jurassic period, but the earliest phase of this evolutionary process remains unclear owing to the exceedingly sparse and spatio-temporally restricted fossil record1-5. Information about the early-diverging species along the avialan line is crucial to understand the evolution of the characteristic bird bauplan, and to reconcile phylogenetic controversies over the origin of birds3,4. Here we describe one of the stratigraphically youngest and geographically southernmost Jurassic avialans, Fujianvenator prodigiosus gen. et sp. nov., from the Tithonian age of China. This specimen exhibits an unusual set of morphological features that are shared with other stem avialans, troodontids and dromaeosaurids, showing the effects of evolutionary mosaicism in deep avialan phylogeny. F. prodigiosus is distinct from all other Mesozoic avialan and non-avialan theropods in having a particularly elongated hindlimb, suggestive of a terrestrial or wading lifestyle-in contrast with other early avialans, which exhibit morphological adaptations to arboreal or aerial environments. During our fieldwork in Zhenghe where F. prodigiosus was found, we discovered a diverse assemblage of vertebrates dominated by aquatic and semi-aquatic species, including teleosts, testudines and choristoderes. Using in situ radioisotopic dating and stratigraphic surveys, we were able to date the fossil-containing horizons in this locality-which we name the Zhenghe Fauna-to 148-150 million years ago. The diversity of the Zhenghe Fauna and its precise chronological framework will provide key insights into terrestrial ecosystems of the Late Jurassic.

New reptile shows dinosaurs and pterosaurs evolved among diverse precursors.

Dinosaurs and pterosaurs have remarkable diversity and disparity through most of the Mesozoic Era1-3. Soon after their origins, these reptiles diversified into a number of long-lived lineages, evolved unprecedented ecologies (for example, flying, large herbivorous forms) and spread across Pangaea4,5. Recent discoveries of dinosaur and pterosaur precursors6-10 demonstrated that these animals were also speciose and widespread, but those precursors have few if any well-preserved skulls, hands and associated skeletons11,12. Here we present a well-preserved partial skeleton (Upper Triassic, Brazil) of the new lagerpetid Venetoraptor gassenae gen. et sp. nov. that offers a more comprehensive look into the skull and ecology of one of these precursors. Its skull has a sharp, raptorial-like beak, preceding that of dinosaurs by around 80 million years, and a large hand with long, trenchant claws that firmly establishes the loss of obligatory quadrupedalism in these precursor lineages. Combining anatomical information of the new species with other dinosaur and pterosaur precursors shows that morphological disparity of precursors resembles that of Triassic pterosaurs and exceeds that of Triassic dinosaurs. Thus, the 'success' of pterosaurs and dinosaurs was a result of differential survival among a broader pool of ecomorphological variation. Our results show that the morphological diversity of ornithodirans started to flourish among early-diverging lineages and not only after the origins of dinosaurs and pterosaurs.

On the first Baryonychinae (Theropoda, Spinosauridae) teeth from South America.

We report the first occurrence in South America of spinosaurid specimens belonging to the clade Baryonychinae. The material comprises three tooth crowns recovered from the Lower Cretaceous (Berriasian-Valanginian) of the Feliz Deserto Formation, State of Sergipe, northeast Brazil. The three specimens have the typical conidont condition, displaying a fluted morphology, veined enamel texture and carinae extending to the cervix region, conditions similar to those shared among spinosaurid theropods. Furthermore, a notable feature is the presence of a serrated carinae composed of 6-7 diminutive denticles per millimetre, a key feature shared by baryonychine spinosaurids. Phylogenetic and multivariate analyses performed on dentition-based and crown-based datasets of theropod taxa confirm the relationship of the three tooth crowns described here with the well-supported spinosaurid clade. Our results retrieve the taxa which comprise the "classically" baryonychine clade as individual branches within a polytomy which includes the three tooth crowns and a subclade of spinosaurines (Spinosaurus + Angaturama/Irritator). Additionally, the discriminant and cluster analyses suggest that the materials are more similar to Suchomimus than to Baryonyx morphometrically, but clearly bearing similarities with baryonychines (sharing a similar morphospace). These new findings paleogeographically expand the occurrence of this clade to western Gondwana, and comprise one of the oldest spinosaurid records worldwide, which brings implications on hypotheses for the emergence and evolution of the clade. Furthermore, these new records indicate the presence of more than one spinosaurid taxon in the Lower Cretaceous deltaic paleoenvironment of the Feliz Deserto Formation.

Theropod dinosaur facial reconstruction and the importance of soft tissues in paleobiology.

Large theropod dinosaurs are often reconstructed with their marginal dentition exposed because of the enormous size of their teeth and their phylogenetic association to crocodylians. We tested this hypothesis using a multiproxy approach. Regressions of skull length and tooth size for a range of theropods and extant varanid lizards confirm that complete coverage of theropod dinosaur teeth with extraoral tissues (gingiva and labial scales) is both plausible and consistent with patterns observed in living ziphodont amniotes. Analyses of dental histology from crocodylians and theropod dinosaurs, including Tyrannosaurus rex, further indicate that the most likely condition was complete coverage of the marginal dentition with extraoral tissue when the mouth was closed. This changes our perceptions about the appearance and oral configuration of these iconic predators and has broad implications for our interpretations of other terrestrial animals with large teeth.

Developmental strategies underlying gigantism and miniaturization in non-avialan theropod dinosaurs.

In amniotes, the predominant developmental strategy underlying body size evolution is thought to be adjustments to the rate of growth rather than its duration. However, most theoretical and experimental studies supporting this axiom focus on pairwise comparisons and/or lack an explicit phylogenetic framework. We present the first large-scale phylogenetic comparative analysis examining developmental strategies underlying the evolution of body size, focusing on non-avialan theropod dinosaurs. We reconstruct ancestral states of growth rate and body mass in a taxonomically rich dataset, finding that contrary to expectations, changes in the rate and duration of growth played nearly equal roles in the evolution of the vast body size disparity present in non-avialan theropods-and perhaps that of amniotes in general.

Iyuku raathi, a new iguanodontian dinosaur from the Early Cretaceous Kirkwood Formation, South Africa.

We name and describe a new iguanodontian dinosaur from the Early Creteceous Kirkwood Formation, Eastern Cape Province, South Africa. This dinosaur is one of only two ornithopod dinosaurs known from the Cretaceous of southern Africa, and is unique in being represented primarily by hatchling to young juvenile individuals as demonstrated by bone histological analysis. All of the juvenile material of this new taxon comes from a single, laterally-restricted bonebed and specimens were primarily recovered as partial to complete single elements, although rare articulated materials and one partial skeleton were found. Sedimentology of the bonebed suggests that this horizon heralds a change in environment upsection to a drier and more seasonal climate. This accumulation of bones is interpreted as seasonal mortality from a nesting site or nesting grounds and may be linked to this environmental shift.

Scleromochlus and the early evolution of Pterosauromorpha.

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.

Low dinosaur biodiversity in central China 2 million years prior to the end-Cretaceous mass extinction.

Whether or not nonavian dinosaur biodiversity declined prior to the end-Cretaceous mass extinction remains controversial as the result of sampling biases in the fossil record, differences in the analytical approaches used, and the rarity of high-precision geochronological dating of dinosaur fossils. Using magnetostratigraphy, cyclostratigraphy, and biostratigraphy, we establish a high-resolution geochronological framework for the fossil-rich Late Cretaceous sedimentary sequence in the Shanyang Basin of central China. We have found only three dinosaurian eggshell taxa (Macroolithus yaotunensis, Elongatoolithus elongatus, and Stromatoolithus pinglingensis) representing two clades (Oviraptoridae and Hadrosauridae) in sediments deposited between ∼68.2 and ∼66.4 million y ago, indicating sustained low dinosaur biodiversity, and that assessment is consistent with the known skeletal remains in the Shanyang and surrounding basins of central China. Along with the dinosaur eggshell records from eastern and southern China, we find a decline in dinosaur biodiversity from the Campanian to the Maastrichtian. Our results support a long-term decline in global dinosaur biodiversity prior to 66 million y ago, which likely set the stage for the end-Cretaceous nonavian dinosaur mass extinction.

A new theropod dinosaur from the early cretaceous (Barremian) of Cabo Espichel, Portugal: Implications for spinosaurid evolution.

Spinosaurids are some of the most enigmatic Mesozoic theropod dinosaurs due to their unique adaptations to aquatic environments and their relative scarcity. Their taxonomy has proven to be especially problematic. Recent discoveries from Western Europe in general, specifically Iberia, provide some of the best specimens for the understanding of their phylogeny, leading to the description of the spinosaurid Vallibonavenatrix cani and the recognition of the Iberian dinosaur Camarillasaurus cirugedae as one of them. Portuguese associated spinosaurid remains (ML1190) from the Papo Seco Formation (early Barremian) were previously assigned to Baryonyx walkeri but new material recovered in 2020 along with new phylogenetic analyses suggests a different phylogenetic placement, making their revision necessary. Here we show that these remains are not attributable to Baryonyx walkeri, but to a new genus and species, Iberospinus natarioi, gen. et sp. nov. The new taxon is characterized by the presence of a single Meckelian foramen in the Meckelian sulcus, a straight profile of the ventral surface of the dentary and a distal thickening of the acromion process of the pubis between other characters. Iberospinus natarioi is recovered as a sister taxon of the clade formed by Baryonyx and Suchomimus, and outside Spinosaurinae when Vallibonaventrix cani is excluded from the analysis. The description of this taxon reinforces Iberia as a hotspot for spinosaur biodiversity, with several endemic taxa for the region. As expected for the clade, the dentary displays a highly vascularized neurovascular network. The morphometric analysis of parts of the skeleton (pedal phalanx and caudal vertebrae, among others) shows an intermediate condition between basal tetanurans and spinosaurines.

Postcranial osteology of Beipiaosaurus inexpectus (Theropoda: Therizinosauria).

Beipiaosaurus inexpectus, from the Lower Cretaceous Yixian Formation (Sihetun locality, near Beipiao), Liaoning, China, is a key taxon for understanding the early evolution of therizinosaurians. Since initial publication in 1999, only the cranial elements of this taxon have been described in detail. Here we present a detailed description of the postcranial skeletal anatomy of the holotype specimen of B. inexpectus, including two never before described dorsal vertebrae from the anterior half of the series. Based on these observations, and comparisons with the postcranial skeleton of therizinosaurian taxa named since the most recent diagnosis, we revised the diagnostic features for B. inexpectus adding three new possible autapomorphies (PII-3 shorter than PIII-4, subequal length of the pre- and postacetabular portions of the ilium, and equidimensional pubic peduncle of ilium). Additionally, we also propose three possible synapomorphies for more inclusive taxa (Therizinosauroidea and Therizinosauridae) and discuss implications for evolutionary trends within Therizinosauria. The newly acquired data from the postcranial osteology of the holotype specimen of B. inexpectus sheds light on our understanding of postcranial skeletal evolution and identification of therizinosaurians.

Quantitative assessment of the vertebral pneumaticity in an anhanguerid pterosaur using micro-CT scanning.

Research on the postcranial skeletal pneumaticity in pterosaurs is common in the literature, but most studies present only qualitative assessments. When quantitative, they are done on isolated bones. Here, we estimate the Air Space Proportion (ASP) obtained from micro-CT scans of the sequence from the sixth cervical to the fourth dorsal vertebra of an anhanguerine pterosaur to understand how pneumaticity is distributed in these bones. Pneumatisation of the vertebrae varied between 68 and 72% of their total volume. The neural arch showed higher ASP in all vertebrae. Anhanguerine vertebral ASP was generally higher than in sauropod vertebrae but lower than in most extant birds. The ASP observed here is lower than that calculated for the appendicular skeleton of other anhanguerian pterosaurs, indicating the potential existence of variation between axial and appendicular pneumatisation. The results point to a pattern in the distribution of the air space, which shows an increase in the area occupied by the trabecular bone in the craniocaudal direction of the vertebral series and, in each vertebra, an increase of the thickness of the trabeculae in the zygapophyses. This indicates that the distribution of pneumatic diverticula in anhanguerine vertebrae may not be associated with stochastic patterns.

Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade.

A remarkably well-preserved, almost complete and articulated new specimen (GP/2E 9266) of Tupandactylus navigans is here described for the Early Cretaceous Crato Formation of Brazil. The new specimen comprises an almost complete skeleton, preserving both the skull and post-cranium, associated with remarkable preservation of soft tissues, which makes it the most complete tapejarid known thus far. CT-Scanning was performed to allow the assessment of bones still covered by sediment. The specimen can be assigned to Tupa. navigans due to its vertical supra-premaxillary bony process and short and rounded parietal crest. It also bears the largest dentary crest among tapejarine pterosaurs and a notarium, which is absent in other representatives of the clade. The new specimen is here regarded as an adult individual. This is the first time that postcranial remains of Tupa. navigans are described, being also an unprecedented record of an articulated tapejarid skeleton from the Araripe Basin.

The first dinosaurs from the Early Cretaceous Hami Pterosaur Fauna, China.

The Early Cretaceous Hami Pterosaur Fauna in Northwest China preserves a large number of specimens of the sexually dimorphic pteranodontoid pterosaur Hamipterus tianshanensis, including 3D eggs and embryos. During the last decade, several more fossils have been collected in this area, including three somphospondylan sauropod specimens. The first is Silutitan sinensis gen. et sp. nov., which consists of an articulated middle to posterior cervical vertebrae series. The second, Hamititan xinjiangensis gen. et sp. nov., consists of an incomplete articulated caudal sequence that could be assigned to lithostrotian titanosaurs based on the strongly procoelous caudal vertebrae with lateral concave surface, as well as marked ventrolateral ridges. The third specimen consists of four sacral vertebral elements, apparently unfused, with exposed camellate internal bone and regarded as somphospondylan. Cladistic analyses based on different datasets recovered Silutitan sinensis as an euhelopodid closely related to Euhelopus and Hamititan xinjiangensis as a titanosaur. Besides the pterosaur Hamipterus and one theropod tooth, these dinosaurs are the first vertebrates reported in this region, increasing the diversity of the fauna as well as the information on Chinese sauropods, further supporting a widespread diversification of somphospondylans during the Early Cretaceous of Asia.

A new styracosternan hadrosauroid (Dinosauria: Ornithischia) from the Early Cretaceous of Portell, Spain.

A new styracosternan ornithopod genus and species is described based on the right dentary of a single specimen from the Mirambell Formation (Early Cretaceous, early Barremian) at the locality of Portell, (Castellón, Spain). Portellsaurus sosbaynati gen. et sp. nov. is diagnosed by two autapomorphic features as well as a unique combination of characters. The autapomorphies include: the absence of a bulge along the ventral margin directly ventral to the base of the coronoid process and the presence of a deep oval cavity on the medial surface of the mandibular adductor fossa below the eleventh-twelfth tooth position. Phylogenetic analyses reveal that the new Iberian form is more closely related to the African taxon Ouranosaurus nigeriensis than to its synchronic Iberian taxa Magnamanus soriaensis and Iguanodon galvensis. In addition, Portellsaurus sosbaynati is less related to other Iberian taxa such as Iguanodon bernissartensis and Proa valdearinnoensis than to the other Early Cretaceous Iberian styracosternans Mantellisaurus atherfieldensis and Morelladon beltrani. A new phylogenetic hypothesis is proposed that resolves Iguanodon (I. bernissartensis, I. galvensis) with the Valanginian Barilium dawsoni into a monophyletic clade (Iguanodontoidea). The recognition of Portellsaurus sosbaynati gen. et sp. nov. as the first styracosternan dinosaur species identified from the Margas de Mirambell Formation (early Barremian-early late Barremian) in the Morella sub-basin (Maestrat Basin, eastern Spain) indicates that the Iberian Peninsula was home to a highly diverse assemblage of medium-to-large bodied styracosternan hadrosauriforms during the Early Cretaceous.

A partial oviraptorosaur skeleton suggests low caenagnathid diversity in the Late Cretaceous Nemegt Formation of Mongolia.

The Nemegt Formation of the Gobi Desert of Mongolia has produced one of the most abundant and diverse oviraptorosaur records globally. However, the caenagnathid component of this fauna remains poorly known. Two caenagnathid taxa are currently recognized from the Nemegt Formation: Elmisaurus rarus and Nomingia gobiensis. Because these taxa are known from mostly non-overlapping material, there are concerns that they could represent the same animal. A partial, weathered caenagnathid skeleton discovered adjacent to the holotype quarry of Nomingia gobiensis is referable to Elmisaurus rarus, revealing more of the morphology of the cranium, mandible, pectoral girdle, and pubis. Despite metatarsals clearly exhibiting autapomorphies of Elmisaurus rarus, overlapping elements are identical to those of Nomingia gobiensis, and add to a growing body of evidence that these taxa represent a single morphotype. In the absence of any positive evidence for two caenagnathid taxa in the Nemegt Formation, Nomingia gobiensis is best regarded as a junior synonym of Elmisaurus rarus. Low caenagnathid diversity in the Nemegt Formation may reflect broader coexistence patterns with other oviraptorosaur families, particularly oviraptorids. In contrast to North America, competition with the exceptionally diverse oviraptorids may have restricted caenagnathids to marginal roles in Late Cretaceous Asian ecosystems.

Dinosaur biodiversity declined well before the asteroid impact, influenced by ecological and environmental pressures.

The question why non-avian dinosaurs went extinct 66 million years ago (Ma) remains unresolved because of the coarseness of the fossil record. A sudden extinction caused by an asteroid is the most accepted hypothesis but it is debated whether dinosaurs were in decline or not before the impact. We analyse the speciation-extinction dynamics for six key dinosaur families, and find a decline across dinosaurs, where diversification shifted to a declining-diversity pattern ~76 Ma. We investigate the influence of ecological and physical factors, and find that the decline of dinosaurs was likely driven by global climate cooling and herbivorous diversity drop. The latter is likely due to hadrosaurs outcompeting other herbivores. We also estimate that extinction risk is related to species age during the decline, suggesting a lack of evolutionary novelty or adaptation to changing environments. These results support an environmentally driven decline of non-avian dinosaurs well before the asteroid impact.

The early origin of a birdlike inner ear and the evolution of dinosaurian movement and vocalization.

Reptiles, including birds, exhibit a range of behaviorally relevant adaptations that are reflected in changes to the structure of the inner ear. These adaptations include the capacity for flight and sensitivity to high-frequency sound. We used three-dimensional morphometric analyses of a large sample of extant and extinct reptiles to investigate inner ear correlates of locomotor ability and hearing acuity. Statistical analyses revealed three vestibular morphotypes, best explained by three locomotor categories-quadrupeds, bipeds and simple fliers (including bipedal nonavialan dinosaurs), and high-maneuverability fliers. Troodontids fall with Archaeopteryx among the extant low-maneuverability fliers. Analyses of cochlear shape revealed a single instance of elongation, on the stem of Archosauria. We suggest that this transformation coincided with the origin of both high-pitched juvenile location, alarm, and hatching-synchronization calls and adult responses to them.

Tooth replacement patterns in the Early Triassic epicynodont Galesaurus planiceps (Therapsida, Cynodontia).

Sixteen specimens of the Early Triassic cynodont Galesaurus planiceps (including eight that were scanned using micro-computed tomography) representing different ontogenetic stages were assembled to study the dental replacement in the species. The growth series shows that the incisors and postcanines continue to develop and replace, even in the largest (presumably oldest) specimen. In contrast, replacement of the canines ceased with the attainment of skeletal maturity, at a basal skull length of ~90 mm, suggesting that Galesaurus had a finite number of canine replacement cycles. Additionally, the functional canine root morphology of these larger specimens showed a tendency to be open-rooted, a condition not previously reported in Mesozoic theriodonts. An alternating pattern of tooth replacement was documented in the maxillary and mandibular postcanine series. Both postcanine series increased in tooth number as the skull lengthened, with the mandibular postcanine series containing more teeth than the maxillary series. In the maxilla, the first postcanine is consistently the smallest tooth, showing a proportional reduction in size as skull length increased. The longer retention of a tooth in this first locus is a key difference between Galesaurus and Thrinaxodon, in which the mesial-most postcanines are lost after replacement. This difference has contributed to the lengthening of the postcanine series in Galesaurus, as teeth continued to be added to the distal end of the tooth row through ontogeny. Overall, there are considerable differences between Galesaurus and Thrinaxodon relating to the replacement and development of their teeth.