Earliest evidence of avian primary feather moult.

Feather moulting is a crucial process in the avian life cycle, which evolved to maintain plumage functionality. However, moulting involves both energetic and functional costs. During moulting, plumage function temporarily decreases between the shedding of old feathers and the full growth of new ones. In flying taxa, a gradual and sequential replacement of flight feathers evolved to maintain aerodynamic capabilities during the moulting period. Little is known about the moult strategies of non-avian pennaraptoran dinosaurs and stem birds, before the emergence of crown lineage. Here, we report on two Early Cretaceous pygostylian birds from the Yixian Formation (125 mya), probably referable to Confuciusornithiformes, exhibiting morphological characteristics that suggest a gradual and sequential moult of wing flight feathers. Short primary feathers interpreted as immature are symmetrically present on both wings, as is typical among extant flying birds. Our survey of the enormous collection of the Tianyu Museum confirms previous findings that evidence of active moult in non-neornithine pennaraptorans is rare and likely indicates a moult cycle greater than one year. Documenting moult in Mesozoic feathered dinosaurs is critical for understanding their ecology, locomotor ability and the evolution of this important life-history process in birds.

Intraspecific variation in the pterosaur Rhamphorhynchus muensteri-implications for flight and socio-sexual signaling.

Pterosaurs were the first powered flying vertebrates, with a fossil record that stretches back to about 230 million years before present. Most species are only known from one to three specimens, which are most often fragmentary. However, Rhamphorhynchus muensteri is known from numerous excellent specimens, including multiple specimens with soft tissue preservation. As such, Rhamphorhynchus muensteri is one of the only pterosaurs amenable to analysis for intraspecific variation. It has been previously predicted that elements directly involved in the flight apparatus, such as those of the forelimb, will be more highly constrained in their proportions than other parts of the skeleton. We investigated the degree of variation seen in elements and body parts of Rhamphorhynchus, which represents the best model system among pterosaurs for testing these expectations of intraspecific variation. We recover evidence for high levels of constraint throughout the appendicular and axial elements (head, neck, torso, tail, forelimbs, hindlimbs), suggesting that all were important for flight. We further find that tail variation increases among the largest specimens, suggesting reduced constraint and/or stronger sexual selection on the tail in more mature individuals.

Unraveling sauropod diversity in the Portezuelo Formation of Patagonia through a comprehensive analysis of new and existing material.

The Portezuelo Formation preserves an outstanding record of the upper Turonian - lower Coniacian. Despite the discovery of a significant quantity of sauropod fossil material from the formation, only two species have been formally described to date: Malarguesaurus florenciae and Futalognkosaurus dukei. Here we present new sauropod material mostly composed of non-articulated caudal vertebrae (MCF-PVPH 916 and 917) that belong to two titanosauriforms on the basis of the following features: anterior caudal vertebrae with procoelous-opisthoplatyan articulations, transverse processes that reach the posterior articular face of the centrum and neural spines with a transverse width of around 50% of their anteroposterior length; anterior and middle caudal vertebrae with the neural arch restricted to the anterior half of the centrum; middle caudal centra with circular cross-section. Phylogenetic analysis recovers the new material in close relation to Malarguesaurus within a monophyletic clade at the base of Somphospondyli. This clade shares large pedicel height with a vertical anterior border on the middle caudal vertebrae, a vertical orientation of the neural spines on the distalmost middle caudal vertebrae and proximalmost posterior caudal vertebrae, and subequal relative lengths of the proximal ulnar condylar processes. The specimens presented here are distinct not only from Futalognkosaurus, but also from other indeterminate titanosaurian remains from the same formation. However, there are no significant differences between the specimen MCF-PVPH 917 and Malarguesaurus, but there are differences between the posterior caudal vertebrae of MCF-PVPH 916 and Malarguesaurus, so they could be considered different species. Whilst we err on the side of caution in not naming new taxa here, the two specimens significantly expand what we know about sauropods in the Turonian-Coniacian ecosystems of Patagonia, which will continue to do so as more material is discovered.

A new stegosaur (Dinosauria: Ornithischia) from the Middle Jurassic of Gansu Province, China.

Stegosaurs are a minor but iconic clade of ornithischian dinosaurs, yet due to a poor fossil record, their early evolution is poorly understood. Here, we describe a new stegosaur, Baiyinosaurus baojiensis, gen. et sp. nov. from the Middle Jurassic Wangjiashan Formation of the Pingchuan District, Baiyin City, Gansu Province, China. The frontal of Baiyinosaurus possesses a unique characteristic among Stegosauria: it is wider than long and contributes to both the medial and anterior margins of the supratemporal fenestra. The character combinations of dorsal vertebrae of Baiyinosaurus are also different to other stegosaurs: its neural arches are not greatly elongated, its parapophyses are well developed, and its neural spines are axially expanded in lateral. The features of the frontal and vertebrae of Baiyinosaurus are reminiscent of basally branching thyreophorans, indicating that Baiyinosaurus is transitional in morphology between early thyreophorans and early-diverging stegosaurs. Systematic analysis shows that Baiyinosaurus is an early-diverging stegosaur.

Possible eucynodont (Synapsida: Cynodontia) tracks from a lacustrine facies in the Lower Jurassic Moenave Formation of southwestern Utah.

Eight fossil tetrapod footprints from lake-shore deposits in the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery Site (SGDS) in southwestern Utah cannot be assigned to the prevalent dinosaurian (Anomoepus, Eubrontes, Gigandipus, Grallator, Kayentapus) or crocodyliform (Batrachopus) ichnotaxa at the site. The tridactyl and tetradactyl footprints are incomplete, consisting of digit- and digit-tip-only imprints. Seven of the eight are likely pes prints; the remaining specimen is a possible manus print. The pes prints have digit imprint morphologies and similar anterior projections and divarication angles to those of Brasilichnium, an ichnotaxon found primarily in eolian paleoenvironments attributed to eucynodont synapsids. Although their incompleteness prevents clear referral to Brasilichnium, the SGDS tracks nevertheless suggest a eucynodont track maker and thus represent a rare, Early Mesozoic occurrence of such tracks outside of an eolian paleoenvironment.

The complexity of tracking stegosaurs and their gregarious behavior.

Stegosaur tracks were unknown until the identification of Deltapodus more than 20 years ago. Currently, the Iberian Peninsula, especially Teruel Province, is one of the areas globally with the most occurrences of these tracks. However, their identification, based on the global record, is problematic due to their similarities with sauropod tracks. A review of the largest number of analyzed Deltapodus tracks globally, including the holotype of D. ibericus and a description of new occurrences, has been carried out. Our research shows substantial morphological variations, but all the studied tracks can be considered D. ibericus based on the manus morphology and the morphometric data. These variations are related to substrate differences and/or different dynamic foot postures (possibly ontogenetically related) during locomotion, as evidenced by changes within the same trackway. We provide detailed comparisons via 3D modeling with sauropod tracks, and our data show that they generally have proportionally longer manus and wider pes because of the differences in the metapodial bones. The scarcity of stegosaur trackways in the fossil record has prevented the identification of gregarious behavior in this group of herbivorous dinosaurs. Two of the studied tracksites show evidence of this behavior, being the only examples among stegosaurs described thus far in the fossil record.

Using linear measurements to diagnose the ecological habitat of Spinosaurus.

Much of the ecological discourse surrounding the polarising theropod Spinosaurus has centred on qualitative discussions. Using a quantitative multivariate data analytical approach on size-adjusted linear measurements of the skull, we examine patterns in skull shape across a range of sauropsid clades and three ecological realms (terrestrial, semi-aquatic, and aquatic). We utilise cluster analyses to identify emergent properties of the data which associate properties of skull shape with ecological realm occupancy. Results revealed terrestrial ecologies to be significantly distinct from both semi- and fully aquatic ecologies, the latter two were not significantly different. Spinosaurids (including Spinosaurus) plotted away from theropods in morphospace and close to both marine taxa and wading birds. The position of nares and the degree of rostral elongation had the greatest effect on categorisation. Comparisons of supervised (k-means) and unsupervised clustering demonstrated categorising taxa into three groups (ecological realms) was inappropriate and suggested instead that cluster division is based on morphological adaptations to feeding on aquatic versus terrestrial food items. The relative position of the nares in longirostrine taxa is associated with which skull bones are elongated. Rostral elongation is observed by either elongating the maxilla and the premaxilla or by elongating the maxilla only. This results in the nares positioned towards the orbits or towards the anterior end of the rostrum respectively, with implications on available feeding methods. Spinosaurids, especially Spinosaurus, show elongation in the maxilla-premaxilla complex, achieving similar functional outcomes to elongation of the premaxilla seen in birds, particularly large-bodied piscivorous taxa. Such a skull construction would bolster "stand-and-wait" predation of aquatic prey to a greater extent than serving other proposed feeding methods.

A new small-sized predatory pseudosuchian archosaur from the Middle-Late Triassic of Southern Brazil.

Before the rise of dinosaurs and pterosaurs, pseudosuchians-reptiles from the crocodilian lineage-dominated the Triassic land ecosystems. This lineage diversified into several less inclusive clades, resulting in a wide ecomorphological diversity during the Middle and Late Triassic. Some giant pseudosuchians occupied the top of the trophic webs, while others developed extensive bony armor as a defense mechanism, which later evolved as a convergence in the avemetatarsalian lineage. On the other hand, there were groups like the Gracilisuchidae, which was composed of carnivorous forms with lightweight build and less than 1 m in length. The fossil record of gracilisuchids is geographically restricted to China and Argentina, with one ambiguous record from Brazil. In the present study, the first unambiguous gracilisuchid from Brazil is described. Parvosuchus aurelioi gen. et sp. nov. comes from the Dinodontosaurus Assemblage Zone of the Santa Maria Formation, which is associated with the Ladinian-Carnian boundary. Composed of a complete cranium, vertebrae, pelvic girdle and hindlimbs, the new species nests with Gracilisuchus stipanicicorum and Maehary bonapartei in a phylogenetic analysis. Its discovery fills a taxonomic gap in Brazilian pseudosuchian fauna and reveals the smallest known member of this clade from the Dinodontosaurus Assemblage Zone, highlighting the diversity of pseudosuchians during the moment that preceded the dawn of dinosaurs.

Diuqin lechiguanae gen. et sp. nov., a new unenlagiine (Theropoda: Paraves) from the Bajo de la Carpa Formation (Neuquén Group, Upper Cretaceous) of Neuquén Province, Patagonia, Argentina.

BACKGROUND: Unenlagiine paravians are among the most relevant Gondwanan theropod dinosaur clades for understanding the origin of birds, yet their fossil record remains incomplete, with most taxa being represented by fragmentary material and/or separated by lengthy temporal gaps, frustrating attempts to characterize unenlagiine evolution. RESULTS AND CONCLUSIONS: Here we describe Diuqin lechiguanae gen. et sp. nov., a new unenlagiine taxon from the Upper Cretaceous (Santonian) Bajo de la Carpa Formation of the Neuquén Basin of Neuquén Province in northern Patagonia, Argentina that fills a substantial stratigraphic gap in the fossil record of these theropods. Although known only from a very incomplete postcranial skeleton, the preserved bones of Diuqin differ from corresponding elements in other unenlagiines, justifying the erection of the new taxon. Moreover, in several morphological aspects, the humerus of Diuqin appears intermediate between those of geologically older unenlagiines from the Neuquén Basin (e.g., Unenlagia spp. from the Turonian-Coniacian Portezuelo Formation) and that of the stratigraphically younger, larger-bodied Austroraptor cabazai from the Campanian-Maastrichtian Allen Formation. Consequently, the morphology of the new taxon appears to indicate a transitional stage in unenlagiine evolution. Phylogenetic analysis recovers Diuqin as a paravian with multiple plausible systematic positions, but the strongest affinity is with Unenlagiinae. The humerus of the new form exhibits subcircular punctures near its distal end that are interpreted as feeding traces most likely left by a conical-toothed crocodyliform, mammal, or theropod, the latter potentially corresponding to a megaraptorid or another unenlagiine individual. Thus, in addition to filling important morphological and temporal gaps in unenlagiine evolutionary history, the new taxon also offers information relating to the paleoecology of these theropods.

Haliskia peterseni, a new anhanguerian pterosaur from the late Early Cretaceous of Australia.

Pterosaur remains have been reported from every continent; however, pterosaur skeletons remain rare. A new pterosaur is presented here, Haliskia peterseni gen. et sp. nov., constituting the most complete specimen from Australia from the upper Albian Toolebuc Formation of the Eromanga Basin (Queensland, Australia). A combination of features, including the presence of a premaxillary crest and curved teeth, and the morphology of the scapulocoracoid, support its referral to Anhangueria. Haliskia can be distinguished from all other anhanguerian pterosaurs based on two dental characters: the 4th and 5th tooth pairs are smaller than the 3rd and 6th, and the 2nd and 5th alveoli are smaller than 3-4 and 6-8. Moreover, the hyoid is 70% the total length of the mandible. The phylogenetic analyses presented here place Haliskia within Anhangueria. In one analysis, Haliskia and Ferrodraco are resolved as sister taxa, with Tropeognathus mesembrinus sister to that clade. The other resolves Haliskia, Mythunga and Ferrodraco in a polytomy within Tropeognathinae. The new Australian pterosaur attests to the success of Anhangueria during the latest Early Cretaceous and suggests that the Australian forms were more taxonomically diverse and palaeobiogeographically complex than previously recognized.

Lokiceratops rangiformis gen. et sp. nov. (Ceratopsidae: Centrosaurinae) from the Campanian Judith River Formation of Montana reveals rapid regional radiations and extreme endemism within centrosaurine dinosaurs.

The Late Cretaceous of western North America supported diverse dinosaur assemblages, though understanding patterns of dinosaur diversity, evolution, and extinction has been historically limited by unequal geographic and temporal sampling. In particular, the existence and extent of faunal endemism along the eastern coastal plain of Laramidia continues to generate debate, and finer scale regional patterns remain elusive. Here, we report a new centrosaurine ceratopsid, Lokiceratops rangiformis, from the lower portion of the McClelland Ferry Member of the Judith River Formation in the Kennedy Coulee region along the Canada-USA border. Dinosaurs from the same small geographic region, and from nearby, stratigraphically equivalent horizons of the lower Oldman Formation in Canada, reveal unprecedented ceratopsid richness, with four sympatric centrosaurine taxa and one chasmosaurine taxon. Phylogenetic results show that Lokiceratops, together with Albertaceratops and Medusaceratops, was part of a clade restricted to a small portion of northern Laramidia approximately 78 million years ago. This group, Albertaceratopsini, was one of multiple centrosaurine clades to undergo geographically restricted radiations, with Nasutuceratopsini restricted to the south and Centrosaurini and Pachyrostra restricted to the north. High regional endemism in centrosaurs is associated with, and may have been driven by, high speciation rates and diversity, with competition between dinosaurs limiting their geographic range. High speciation rates may in turn have been driven in part by sexual selection or latitudinally uneven climatic and floral gradients. The high endemism seen in centrosaurines and other dinosaurs implies that dinosaur diversity is underestimated and contrasts with the large geographic ranges seen in most extant mammalian megafauna.

The last ceratosaur of Asia: a new noasaurid from the Early Cretaceous Great Siberian Refugium.

The noasaurid ceratosaur Kiyacursor longipes gen. et sp. nov. is described based on a fragmentary skeleton including cervical vertebra, pectoral girdle, humerus and hind limbs from the Lower Cretaceous (Aptian) Ilek Formation at Shestakovo 1 locality in Western Siberia, Russia. This is the first ceratosaur from the Early Cretaceous of Asia, extending the stratigraphic range of Ceratosauria by 40 Myr on that continent. Kiyacursor shares unique hind limb proportions with Elaphrosaurus and Limusaurus, suggesting improved cursorial ability. These taxa show an ostrich-like specialization of the pes, with a large third metatarsal and greatly reduced second metatarsal. By contrast, all other fast running non-avian theropod dinosaurs have an arctometatarsalian pes, with the third metatarsal strongly reduced proximally. The new taxon lived in the Early Cretaceous ecosystem containing a number of other Jurassic relics, such as stem salamanders, protosuchian and shartegosuchid crocodyliforms, tritylodontid synapsids and docodontan mammaliaforms.

A vertebra of a small species of Pachycetus from the North Sea and its inner structure and vascularity compared with other basilosaurid vertebrae from the same site.

In the Western Scheldt Estuary near the Belgian-Dutch border, middle to late Eocene strata crop out at the current seafloor. Most vertebrae of large Eocene basilosaurid taxa from this area were previously described in several papers. They represent three morphotypes: elongated vertebrae of a large species of Pachycetus (Morphotype 1b), a not-elongated vertebra of a large 'dorudontid' basilosaurid (Morphotype 2) and 'shortened' vertebrae of a new, unnamed taxon (Morphotype 3). This article deals with a still undescribed, smaller vertebra, NMR-16642, from this site. Our first aim was to date it by dinoflagellate cysts in adhering sediments. Yielding an age of about 38 Ma, it is one of the very few remains of basilosaurids from Europe, of which the age could be assessed with reasonable certainty. The vertebra, Morphotype 1a, is assigned to a small species of Pachycetus. High-quality CT scans are used to differentiate between NMR-16642, Morphotype 1a, and the large species of Pachycetus, Morphotype 1b. Another aim of this paper is to investigate the inner structure and vascularity of the study vertebra and that of the other morphotypes (1b, 2, 3) from this area by using high-quality CT scans. Notwithstanding differences in size, shape and compactness, the vertebral inner structure with a multi-layered cortex of periosteal bone, surrounding two cones of endosteal bone appears to be basically similar in all morphotypes. Apparently, this inner structure reflects the ontogenetic vertebral growth. An attempt to reconstruct the vascularity of the vertebrae reveals a remarkable pattern of interconnected vascular systems. From the dorsal and, if present, ventral foramina, vascular canals are running to a central vascular node. From this node a system of vascular canals goes to the epiphyseal ends, giving rise to separate systems for cortex and cones. It is the first time that the vascularity of vertebrae of archaeocetes is investigated.

A new abelisaurid dinosaur from the end Cretaceous of Patagonia and evolutionary rates among the Ceratosauria.

Gondwanan dinosaur faunae during the 20 Myr preceding the Cretaceous-Palaeogene (K/Pg) extinction included several lineages that were absent or poorly represented in Laurasian landmasses. Among these, the South American fossil record contains diverse abelisaurids, arguably the most successful groups of carnivorous dinosaurs from Gondwana in the Cretaceous, reaching their highest diversity towards the end of this period. Here we describe Koleken inakayali gen. et sp. n., a new abelisaurid from the La Colonia Formation (Maastrichtian, Upper Cretaceous) of Patagonia. Koleken inakayali is known from several skull bones, an almost complete dorsal series, complete sacrum, several caudal vertebrae, pelvic girdle and almost complete hind limbs. The new abelisaurid shows a unique set of features in the skull and several anatomical differences from Carnotaurus sastrei (the only other abelisaurid known from the La Colonia Formation). Koleken inakayali is retrieved as a brachyrostran abelisaurid, clustered with other South American abelisaurids from the latest Cretaceous (Campanian-Maastrichtian), such as Aucasaurus, Niebla and Carnotaurus. Leveraging our phylogeny estimates, we explore rates of morphological evolution across ceratosaurian lineages, finding them to be particularly high for elaphrosaurine noasaurids and around the base of Abelisauridae, before the Early Cretaceous radiation of the latter clade. The Noasauridae and their sister clade show contrasting patterns of morphological evolution, with noasaurids undergoing an early phase of accelerated evolution of the axial and hind limb skeleton in the Jurassic, and the abelisaurids exhibiting sustained high rates of cranial evolution during the Early Cretaceous. These results provide much needed context for the evolutionary dynamics of ceratosaurian theropods, contributing to broader understanding of macroevolutionary patterns across dinosaurs.

Early Jurassic origin of avian endothermy and thermophysiological diversity in dinosaurs.

A fundamental question in dinosaur evolution is how they adapted to long-term climatic shifts during the Mesozoic and when they developed environmentally independent, avian-style acclimatization, becoming endothermic.1,2 The ability of warm-blooded dinosaurs to flourish in harsher environments, including cold, high-latitude regions,3,4 raises intriguing questions about the origins of key innovations shared with modern birds,5,6 indicating that the development of homeothermy (keeping constant body temperature) and endothermy (generating body heat) played a crucial role in their ecological diversification.7 Despite substantial evidence across scientific disciplines (anatomy,8 reproduction,9 energetics,10 biomechanics,10 osteohistology,11 palaeobiogeography,12 geochemistry,13,14 and soft tissues15,16,17), a consensus on dinosaur thermophysiology remains elusive.1,12,15,17,18,19 Differential thermophysiological strategies among terrestrial tetrapods allow endotherms (birds and mammals) to expand their latitudinal range (from the tropics to polar regions), owing to their reduced reliance on environmental temperature.20 By contrast, most reptilian lineages (squamates, turtles, and crocodilians) and amphibians are predominantly constrained by temperature in regions closer to the tropics.21 Determining when this macroecological pattern emerged in the avian lineage relies heavily on identifying the origin of these key physiological traits. Combining fossils with macroevolutionary and palaeoclimatic models, we unveil distinct evolutionary pathways in the main dinosaur lineages: ornithischians and theropods diversified across broader climatic landscapes, trending toward cooler niches. An Early Jurassic shift to colder climates in Theropoda suggests an early adoption of endothermy. Conversely, sauropodomorphs exhibited prolonged climatic conservatism associated with higher thermal conditions, emphasizing temperature, rather than plant productivity, as the primary driver of this pattern, suggesting poikilothermy with a stronger dependence on higher temperatures in sauropods.

Using your head - cranial steering in pterosaurs.

The vast majority of pterosaurs are characterized by relatively large, elongate heads that are often adorned with large, elaborate crests. Projecting out in front of the body, these large heads and any crests must have had an aerodynamic effect. The working hypothesis of the present study is that these oversized heads were used to control the left-right motions of the body during flight. Using digital models of eight non-pterodactyloids ("rhamphorhyncoids") and ten pterodactyloids, the turning moments associated with the head + neck show a close and consistent correspondence with the rotational inertia of the whole body about a vertical axis in both groups, supporting the idea of a functional relationship. Turning moments come from calculating the lateral area of the head (plus any crests) and determining the associated lift (aerodynamic force) as a function of flight speed, with flight speeds being based on body mass. Rotational inertias were calculated from the three-dimensional mass distribution of the axial body, the limbs, and the flight membranes. The close correlation between turning moment and rotational inertia was used to revise the life restorations of two pterosaurs and to infer relatively lower flight speeds in another two.

Cellular structure of dinosaur scales reveals retention of reptile-type skin during the evolutionary transition to feathers.

Fossil feathers have transformed our understanding of integumentary evolution in vertebrates. The evolution of feathers is associated with novel skin ultrastructures, but the fossil record of these changes is poor and thus the critical transition from scaled to feathered skin is poorly understood. Here we shed light on this issue using preserved skin in the non-avian feathered dinosaur Psittacosaurus. Skin in the non-feathered, scaled torso is three-dimensionally replicated in silica and preserves epidermal layers, corneocytes and melanosomes. The morphology of the preserved stratum corneum is consistent with an original composition rich in corneous beta proteins, rather than (alpha-) keratins as in the feathered skin of birds. The stratum corneum is relatively thin in the ventral torso compared to extant quadrupedal reptiles, reflecting a reduced demand for mechanical protection in an elevated bipedal stance. The distribution of the melanosomes in the fossil skin is consistent with melanin-based colouration in extant crocodilians. Collectively, the fossil evidence supports partitioning of skin development in Psittacosaurus: a reptile-type condition in non-feathered regions and an avian-like condition in feathered regions. Retention of reptile-type skin in non-feathered regions would have ensured essential skin functions during the early, experimental stages of feather evolution.