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Alvarezsauria is a group of morphologically distinctive, medium- to small-sized later-diverging coelurosaurian theropod dinosaurs, whose record ranges from the Late Jurassic to the Late Cretaceous. This clade had a widespread distribution in Laurasia in what is now Europe, Asia, and North America, although there are also several Cretaceous taxa from Gondwana in what is now Argentina that all belong to the family Alvarezsauridae. Although alvarezsaurid taxonomic diversity and anatomical knowledge has expanded over the last decade, alvarezsaurid internal phylogenetic relationships remain highly debated. In this contribution, we describe new alvarezsaurid material and review previously reported alvarezsaurid specimens from the middle Campanian-lower Maastrichtian Allen Formation discovered at the Salitral Ojo de Agua locality in Río Negro province, Patagonia, Argentina. Although these specimens were collected at different times without precise original provenance information, here we gathered X-ray diffraction data of the associated sediments that now suggests that all of these specimens come from the same site and stratigraphic level as the holotype of the alvarezsaurid Bonapartenykus ultimus. Based on this new provenance information and the morphological similarity of the considered specimens, here we tentatively refer them to the genus Bonapartenykus. These newly referred specimens add to the record of this genus and shed light on the body plan of the alvarezsaurid clade Patagonykinae by permitting a more complete reconstruction of the neck, pectoral girdle, hindlimb, and tail.

Alvarezsauria is a group of morphologically distinctive, medium- to small-sized later-diverging coelurosaurian theropod dinosaurs, whose record ranges from the Late Jurassic to the Late Cretaceous. This clade had a widespread distribution in Laurasia in what is now Europe, Asia, and North America, although there are also several Cretaceous taxa from Gondwana in what is now Argentina that all belong to the family Alvarezsauridae. Although alvarezsaurid taxonomic diversity and anatomical knowledge has expanded over the last decade, alvarezsaurid internal phylogenetic relationships remain highly debated. In this contribution, we describe new alvarezsaurid material and review previously reported alvarezsaurid specimens from the middle Campanian-lower Maastrichtian Allen Formation discovered at the Salitral Ojo de Agua locality in Río Negro province, Patagonia, Argentina. Although these specimens were collected at different times without precise original provenance information, here we gathered X-ray diffraction data of the associated sediments that now suggests that all of these specimens come from the same site and stratigraphic level as the holotype of the alvarezsaurid Bonapartenykus ultimus. Based on this new provenance information and the morphological similarity of the considered specimens, here we tentatively refer them to the genus Bonapartenykus. These newly referred specimens add to the record of this genus and shed light on the body plan of the alvarezsaurid clade Patagonykinae by permitting a more complete reconstruction of the neck, pectoral girdle, hindlimb, and tail.

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.

Dilophosaurus wetherilli was the largest animal known to have lived on land in North America during the Early Jurassic. Despite its charismatic presence in pop culture and dinosaurian phylogenetic analyses, major aspects of the skeletal anatomy, taxonomy, ontogeny, and evolutionary relationships of this dinosaur remain unknown. Skeletons of this species were collected from the middle and lower part of the Kayenta Formation in the Navajo Nation in northern Arizona. Redescription of the holotype, referred, and previously undescribed specimens of Dilophosaurus wetherilli supports the existence of a single species of crested, large-bodied theropod in the Kayenta Formation. The parasagittal nasolacrimal crests are uniquely constructed by a small ridge on the nasal process of the premaxilla, dorsoventrally expanded nasal, and tall lacrimal that includes a posterior process behind the eye. The cervical vertebrae exhibit serial variation within the posterior centrodiapophyseal lamina, which bifurcates and reunites down the neck. Iterative specimen-based phylogenetic analyses result in each of the additional specimens recovered as the sister taxon to the holotype. When all five specimens are included in an analysis, they form a monophyletic clade that supports the monotypy of the genus. Dilophosaurus wetherilli is not recovered as a ceratosaur or coelophysoid, but is instead a non-averostran neotheropod in a grade with other stem-averostrans such as Cryolophosaurus ellioti and Zupaysaurus rougieri. We did not recover a monophyletic ‘Dilophosauridae.’ Instead of being apomorphic for a small clade of early theropods, it is more likely that elaboration of the nasals and lacrimals of stem-averostrans is plesiomorphically present in early ceratosaurs and tetanurans that share those features. Many characters of the axial skeleton of Dilophosaurus wetherilli are derived compared to Late Triassic theropods and may be associated with macropredation and an increase in body size in Theropoda across the Triassic-Jurassic boundary.

In recent decades, intensive research on non-avian dinosaurs has strongly suggested that these animals were restricted to terrestrial environments 1 . Historical proposals that some groups, such as sauropods and hadrosaurs, lived in aquatic environments 2 , 3 were abandoned decades ago 4 , 5 – 6 . It has recently been argued that at least some of the spinosaurids—an unusual group of large-bodied theropods of the Cretaceous era—were semi-aquatic 7 , 8 , but this idea has been challenged on anatomical, biomechanical and taphonomic grounds, and remains controversial 9 , 10 – 11 . Here we present unambiguous evidence for an aquatic propulsive structure in a dinosaur, the giant theropod Spinosaurus aegyptiacus 7 , 12 . This dinosaur has a tail with an unexpected and unique shape that consists of extremely tall neural spines and elongate chevrons, which forms a large, flexible fin-like organ capable of extensive lateral excursion. Using a robotic flapping apparatus to measure undulatory forces in physical models of different tail shapes, we show that the tail shape of Spinosaurus produces greater thrust and efficiency in water than the tail shapes of terrestrial dinosaurs and that these measures of performance are more comparable to those of extant aquatic vertebrates that use vertically expanded tails to generate forward propulsion while swimming. These results are consistent with the suite of adaptations for an aquatic lifestyle and piscivorous diet that have previously been documented for Spinosaurus 7 , 13 , 14 . Although developed to a lesser degree, aquatic adaptations are also found in other members of the spinosaurid clade 15 , 16 , which had a near-global distribution and a stratigraphic range of more than 50 million years 14 , pointing to a substantial invasion of aquatic environments by dinosaurs. Discovery that the giant theropod dinosaur Spinosaurus has a large flexible tail indicates that it was primarily aquatic and swam in a similar manner to extant tail-propelled aquatic vertebrates.

A recently discovered fossil belonging to the Scansoriopterygidae, a group of bizarre dinosaurs closely related to birds, represents a new scansoriopterygid species and preserves evidence of a membranous aerodynamic surface very different from a classic avian wing. Taking flight by the wrist At the base of the dinosaur lineage that eventually led to birds there is a small group of bizarre dinosaurs called scansoriopterygids. They were very small, often with long digits and are usually reconstructed as tree-dwelling lemur-like creatures. Here Xing Xu and colleagues present what is possibly the strangest scansoriopterygid so far discovered. Named Yi qi , the tiny animal, from 160 million-year-old sediments in China, has an unusual assortment of stiff filamentous feathers and also two long bony elements attached to the wrists, unlike anything seen previously in any dinosaur. These structures, resembling extraneous bones seen in a variety of tetrapods, may have supported a membrane that might have sustained gliding flight. Traces of such a membrane are preserved with the specimen. The wings of birds and their closest theropod relatives share a uniform fundamental architecture, with pinnate flight feathers as the key component 1 , 2 , 3 . Here we report a new scansoriopterygid theropod, Yi qi gen. et sp. nov., based on a new specimen from the Middle–Upper Jurassic period Tiaojishan Formation of Hebei Province, China 4 . Yi is nested phylogenetically among winged theropods but has large stiff filamentous feathers of an unusual type on both the forelimb and hindlimb. However, the filamentous feathers of Yi resemble pinnate feathers in bearing morphologically diverse melanosomes 5 . Most surprisingly, Yi has a long rod-like bone extending from each wrist, and patches of membranous tissue preserved between the rod-like bones and the manual digits. Analogous features are unknown in any dinosaur but occur in various flying and gliding tetrapods 6 , 7 , 8 , 9 , 10 , suggesting the intriguing possibility that Yi had membranous aerodynamic surfaces totally different from the archetypal feathered wings of birds and their closest relatives. Documentation of the unique forelimbs of Yi greatly increases the morphological disparity known to exist among dinosaurs, and highlights the extraordinary breadth and richness of the evolutionary experimentation that took place close to the origin of birds.

Among extant vertebrates, pneumatization of postcranial bones is unique to birds, with few known exceptions in other groups. Through reduction in bone mass, this feature is thought to benefit flight capacity in modern birds, but its prevalence in non-avian dinosaurs of variable sizes has generated competing hypotheses on the initial adaptive significance of postcranial pneumaticity. To better understand the evolutionary history of postcranial pneumaticity, studies have surveyed its distribution among non-avian dinosaurs. Nevertheless, the degree of pneumaticity in the basal coelurosaurian group Ornithomimosauria remains poorly known, despite their potential to greatly enhance our understanding of the early evolution of pneumatic bones along the lineage leading to birds. Historically, the identification of postcranial pneumaticity in non-avian dinosaurs has been based on examination of external morphology, and few studies thus far have focused on the internal architecture of pneumatic structures inside the bones. Here, we describe the vertebral pneumaticity of the ornithomimosaur Archaeornithomimus with the aid of X-ray computed tomography (CT) imaging. Complementary examination of external and internal osteology reveals (1) highly pneumatized cervical vertebrae with an elaborate configuration of interconnected chambers within the neural arch and the centrum; (2) anterior dorsal vertebrae with pneumatic chambers inside the neural arch; (3) apneumatic sacral vertebrae; and (4) a subset of proximal caudal vertebrae with limited pneumatic invasion into the neural arch. Comparisons with other theropod dinosaurs suggest that ornithomimosaurs primitively exhibited a plesiomorphic theropod condition for axial pneumaticity that was extended among later taxa, such as Archaeornithomimus and large bodied Deinocheirus. This finding corroborates the notion that evolutionary increases in vertebral pneumaticity occurred in parallel among independent lineages of bird-line archosaurs. Beyond providing a comprehensive view of vertebral pneumaticity in a non-avian coelurosaur, this study demonstrates the utility and need of CT imaging for further clarifying the early evolutionary history of postcranial pneumaticity.

Powered flight evolved independently in vertebrates in the pterosaurs, birds and bats, each of which has a different configuration of the bony elements and epidermal structures that form the wings 1 , 2 . Whereas the early fossil records of pterosaurs and bats are sparse, mounting evidence (primarily from China) of feathered non-avian dinosaurs and stemward avians that derive primarily from the Middle–Upper Jurassic and Lower Cretaceous periods has enabled the slow piecing together of the origins of avian flight 3 , 4 . These fossils demonstrate that, close to the origin of flight, dinosaurs closely related to birds were experimenting with a diversity of wing structures 3 , 5 . One of the most surprising of these is that of the scansoriopterygid (Theropoda, Maniraptora) Yi qi , which has membranous wings—a flight apparatus that was previously unknown among theropods but that is used by both the pterosaur and bat lineages 6 . This observation was not universally accepted 7 . Here we describe a newly identified scansoriopterygid—which we name Ambopteryx longibrachium , gen. et sp. nov.—from the Upper Jurassic period. This specimen provides support for the widespread existence of membranous wings and the styliform element in the Scansoriopterygidae, as well as evidence for the diet of this enigmatic theropod clade. Our analyses show that marked changes in wing architecture evolved near the split between the Scansoriopterygidae and the avian lineage, as the two clades travelled along very different paths to becoming volant. The membranous wings supported by elongate forelimbs that are present in scansoriopterygids probably represent a short-lived experimentation with volant behaviour, and feathered wings were ultimately favoured during the later evolution of Paraves. A newly described Jurassic scansoriopterygid reveals that membranous wings and the styliform element were widespread in this clade, which suggests that a distinctive wing architecture evolved near the split between the Scansoriopterygidae and the paravian lineages.