Search

Ornithopod dinosaurs appeared during the Middle Jurassic, but it was in the Lower Cretaceous they started their successful evolutionary history. Different phylogenies describing the evolutionary relationships of Ornithopoda are mostly based on cranial features, however there is a lack of well-preserved and complete skulls for the basal member of the clade, hampering our knowledge on the mode and tempo of these herbivorous dinosaurs. Here we describe YLSNHM 01942, a well-preserved skull of a juvenile neornithischian from the Liaoning Province of China. The specimen was scanned with a μCT scan, and all the elements were segmented and extrapolated for description. The specimen shows a ventral deformation due to the compression of the sediment, and a few rostral elements were artificially added. The specimen is attributed to the basal ornithopod Jeholosaurus shangyuanensis because of the presence of a large foramen in the quadratojugal, however it lacks the nodular ornamentation on the postorbital and jugal, herein interpreted as an ontogenetic feature. This, together with the disarticulation degree of the cranial elements, suggest YLSNHM 01942 represents a juvenile Jeholosaurus. The endosseous labyrinth is tentatively reconstructed, although the disarticulation of the neurocranial bones hampers its complete reconstruction. Thanks to the analysis of previously undescribed inner neurocranial bones (such as the prootics, the exoccipital/ophistotic, basioccipital, and basisphenoid), we improve the previous phylogenetical scoring for J. shangyuanensis, and perform a phylogenetical analysis adding the basal ornithopod Changmiania liaoningensis and the recently re-evaluated Ajkaceratops kozmai. The phylogenetical analysis reports a well-supported base of Ornithopoda, with C. liaoningensis as the most basal ornithopod, and a resolved topology for Nanosaurus agilis, Changchunsaurus parvus, Haya griva, Yandusaurus hongheensis and J. shangyuanensis., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2025 Bertozzo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Pterosaurs were the first vertebrates to evolve active flight. The lack of many well-preserved pterosaur fossils limits our understanding of the functional anatomy and behavior of these flight pioneers, particularly from their early history (Triassic to Middle Jurassic). Here we describe in detail the osteology of an exceptionally preserved Middle Jurassic pterosaur, the holotype of Dearc sgiathanach from the Isle of Skye, Scotland. We identify new autapomorphies of the flight apparatus (humerus and sternum), which further support the distinctiveness of Dearc compared with other early-diverging pterosaurs and describe features, such as the vertebral morphology, shared with later-diverging pterosaurs that probably developed convergently to support a large body size or as a sign of modular evolution. We used extant phylogenetic bracketing to infer the principal cranial and antebrachial musculature, indicating that Dearc had large and anteriorly placed palatal musculature that compensated for weak temporal jaw adductors and wing musculature suggestive of flight style reliant on powerful adduction and protraction of the humerus. Comparisons with other pterosaurs revealed that non-pterodactyloids such as Dearc, despite their overall conservative bauplans, adapted various flight and feeding styles. The osteology and myology of Dearc are indicative of a large predator that flew and hunted above lagoons and nearshore environments of the Middle Jurassic., Competing Interests: Declarations. Ethics approval and consent to participate: The specimen was excavated from a Site of Special Scientific Interest (SSSIs) in Scotland, under permission from the Scotland’s Nature Agency. The holotype specimen is stored in the National Museum of Scotland (Edinburgh, Scotland, UK) under the collection number NMS G.2021.6.1–4. This is a public repository where the specimen is permanently available for research. Consent for publication: All authors approved the text. Competing interests: The authors declare no competing interests., (© 2025. Crown.)

The first partial skeleton of a carcharodontosaurid theropod was described from the Egyptian Bahariya Oasis by Ernst Stromer in 1931. Stromer referred the specimen to the species Megalosaurus saharicus, originally described on the basis of isolated teeth from slightly older rocks in Algeria, under the new genus name Carcharodontosaurus saharicus. Unfortunately, almost all of the material from the Bahariya Oasis, including the specimen of Carcharodontosaurus was destroyed during World War II. In 1996, a relatively complete carcharodontosaurid cranium was described from similar aged rocks in Morocco and designated the neotype of the species Carcharodontosaurus saharicus in 2007. However, due to the destruction of the original material, comparisons of the neotype to the Egyptian fossils have so far only been done cursorily. A detailed reexamination of the available information on the Egyptian carcharodontosaurid, including a previously undescribed photograph of the exhibited specimen, reveals that it differs from the Moroccan neotype in numerous characters, such as the development of the emargination of the antorbital fossa on the nasals, the presence of a horn-like rugosity on the nasal, the lack of a dorsoventral expansion of the lacrimal contact on the frontals, and the relative enlargement of the cerebrum. The referability of the Egyptian specimen to the Algerian M. saharicus is found to be questionable, and the neotype designation of the Moroccan material for C. saharicus is accepted here under consideration of ICZN Atricle 75, as it both compares more favorably to M. saharicus and originates from a locality closer to the type locality. A new genus and species, Tameryraptor markgrafi gen. et sp. nov, is proposed for the Egyptian taxon. The theropods of the Bahariya Oasis and the Moroccan Kem Kem Group are thus not as closely related as previously thought, and the proposed faunal similarities between these two strata need further examination., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2025 Kellermann et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Rhamphorhynchus is one of the best-known pterosaurs, with well over 100 specimens being held in public collections. Most of these represent juvenile animals, and the adults known are typically around 1 m in wingspan. Here we describe a near complete skeleton, preserved partially in 3D, of an animal with a wingspan of around 1.8 m, that is considerably larger than other known specimens, and is among the largest known non-pterodactyloid pterosaurs. This animal shows differences in the anatomy not seen in smaller specimens, revealing details of late-stage ontogeny in this genus. The specimen exhibits a disproportionate reduction in the size of the orbit and increase in the size of the lower temporal fenestra, a reduction in the proportional mandibular symphysis, and unusually laterally flattened teeth, which may point to a changing diet as these animals grew. These features show a transition from smaller to larger specimens of Rhamphorhynchus and also appear in other large specimens of rhamphorhyhchines and point to a consistent pattern in their development., Competing Interests: David W.E. Hone is an Academic Editor for PeerJ., (©2025 Hone and McDavid.)

Among the most revolutionary insights emerging from 200 years of research on dinosaurs is that the clade Dinosauria is represented by approximately 11 000 living species of birds. Although the origin of birds among dinosaurs has been reviewed extensively, recent years have witnessed tremendous progress in our understanding of the deep evolutionary origins of numerous distinctive avian anatomical systems. These advances have been enabled by exciting new fossil discoveries, leading to an ever-expanding phylogenetic framework with which to pinpoint the origins of characteristic avian features. The present review focuses on four notable avian systems whose Mesozoic evolutionary history has been greatly clarified by recent discoveries: brain, kinetic palate, pectoral girdle and postcranial skeletal pneumaticity.

Since the start of the twenty-first century, there has been a notable increase in annual publications focusing on dinosaur reproduction and ontogeny with researchers using these data to address a range of macroevolutionary questions about dinosaurs. Ontogeny, which is closely tied to osteological morphological variation, impacts several key research areas, such as taxonomic diversity, population dynamics, palaeoecology, macroevolution, as well as the physiological and reproductive factors driving ecological success. While these broad studies have significantly advanced our understanding of dinosaur evolution, they have also revealed important challenges and areas needing further investigation. In this review, we aim to outline some of these challenges in major research areas linked to dinosaur ontogeny, namely reproductive biology, osteohistological growth strategies, morphological osteological variation and the link between ontogeny and macroevolution. We also offer some recommendations for best practices and promising future research directions. These recommendations include increasing sample sizes through fieldwork and exhaustive use of pre-existing fossil collections, using micro-computed tomography (μCT) scanning methods to increase dataset sizes in a non-destructive manner, methodical collection and reposition of μCT scan data, assessing ontogenetic maturity, establishing consistency in terminology and methods and building comprehensive extant comparative datasets.

Dinosaur locomotor biomechanics are of major interest. Locomotion of an animal affects many, if not most, aspects of life reconstruction, including behaviour, performance, ecology and appearance. Yet locomotion is one aspect of non-avian dinosaurs that we cannot directly observe. To shed light on how dinosaurs moved, we must draw from multiple sources of evidence. Extant taxa provide the basic principles of locomotion, bracket soft-tissue reconstructions and provide validation data for methods and hypotheses applied to dinosaurs. The skeletal evidence itself can be used directly to reconstruct posture, range of motion and mass (segment and whole-body). Building on skeletal reconstructions, musculoskeletal models inform muscle function and form the basis of simulations to test hypotheses of locomotor performance. Finally, fossilized footprints are our only direct record of motion and can provide important snapshots of extinct animals, shedding light on speed, gait and posture. Building confident reconstructions of dinosaur locomotion requires evidence from all four sources of information. This review explores recent work in these areas, with a methodological focus.

Biomechanical studies in sauropod dinosaurs are mainly focused on neck posture and feeding strategy. Few works investigate other aspects such as tail movement and function, especially in the clade Titanosauria, the most diverse within Neosauropoda. This study applied biomechanical concepts of neutral pose (cartilaginous neutral pose) and range of motion to verify the shape and direction of the caudal region of the advanced titanosaurs Adamantisaurus mezzalirai and Baurutitan britoi, in addition to comparing it with other titanosaurs (e.g., Arrudatitan maximus, Lirainosaurus astibiae, and Trigonosaurus pricei). As a result, the tails analyzed have a sigmoidal-convex shape, probably close to the ground (but not touching it). The sigmoidal-convex shape could increase the moment arm for the M. caudofemoralis longus, making it possible to use the tail as a fifth stabilizing member. This work expands our knowledge about sauropod dinosaurs by exposing a diversity of shapes for the tails of titanosaur sauropods and brings new possibilities for how these dinosaurs could use their tails., (© 2024 Anatomical Society.)

We present the pelvic and hindlimb musculature of the abelisaurid Skorpiovenator bustingorryi, constituting the most comprehensive muscle reconstruction to date in ceratosaur theropods. Using extant phylogenetic bracket method, we reconstructed 39 muscles that can commonly found in extant archosaurs. Through the identification of bone correlates, we recognized thigh and hindlimb muscles including knee extensors, m. iliofibularis, m. flexor tibialis externus, mm. caudofemorales, mm. puboischiofemorales, and crus muscles important in foot extension and flexion (e.g., m. tibialis anterior, mm. gastrocnemii). Also, autopodial intrinsic muscles were reconstructed whose function involve extension (m. extensor digiti 2-4), flexion (mm. flexor digitorum brevis superficialis), interdigital adduction (m. interosseus dorsalis) and abduction (m. interosseous plantaris, m. abductor 4). Abelisaurids like Skorpiovenator show a deep pre- and postacetabular blade of the ilia and enlarged cnemial crests, which would have helped increasing the moment arm of muscles related to hip flexion and hindlimb extension. Also, pedal muscles related to pronation were probably present but reduced (e.g., m. pronator profundus). Despite some gross differences in the autopodial morphology in extant outgroups (e.g., crocodilian metatarsus and avian tarsometatarsus), the present study allows us to hypothesize several pedal muscles in Skorpiovenator. These muscles would not be arranged in tendinous bundles as in Neornithes, but rather the condition would be similar to that of crocodilians with several layers formed by fleshy bellies on the plantar and dorsal aspects of the metatarsus. The musculature of Skorpiovenator is key for future studies concerning abelisaurid biomechanics, including the integration of functional morphology and ichnological data., (© 2024 American Association for Anatomy.)

Since Hampé's classic developmental experiments in the mid-twentieth century 1,2 , the reduced avian fibula has sparked sustained curiosity 3-6 . The fibula transformed throughout dinosaur evolution from a columnar structure into its splint-like avian form, a change long thought to be of little biomechanical consequence 3,6 . Here we integrated comparative three-dimensional kinematic analyses with transitional morphologies from the fossil record to refute this assumption and show that the reduced fibula serves a crucial function in enabling extreme knee long-axis rotation (LAR). Extreme LAR is fundamental to avian locomotion and is regularly exploited by living birds to execute complex terrestrial manoeuvres 7 . We infer that the evolution of this capacity was preceded by restriction of the knee to hinge-like motion in early theropod dinosaurs, driven by the origin of a mid-shank articulation 8 that precluded ancestral patterns of tibiofibular motion. Freeing of the fibula from the ankle joint later enabled mobilization of this initially static articulation and, in doing so, established a novel pattern of tibiofibular kinematics essential to the extreme levels of LAR retained by modern birds. Fibular reduction thus ushered in a transition to LAR-dominated three-dimensional limb control, profoundly altering the course of theropod locomotor evolution., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Well-preserved specimens of a new species of arthrodiran placoderm, Groenlandaspis howittensis sp. nov. (Middle Devonian of Victoria, Australia), reveals previously unknown information on the dermal skeleton, body-shape and dentition of the wide-spread genus Groenlandaspis . The new material includes dual pineal plates, extrascapular plates, and cheek bones showing the presence of cutaneous sensory organs. The anterior supragnathal, usually a paired element in arthrodires, is a fused median bone in G. howittensis sp. nov. It is positioned anterior to the occlusion of the mouth between the lower jaw (infragnathals) and upper jaw (posterior supragnathals) bones, indicating a specialised feeding mechanism and broadening the known diversity of placoderm dental morphologies. G. howittensis sp. nov. differs from all other groenlandaspidids by a less pronounced posterior expansion of the nuchal plate; the shape of the posterior dorsolateral plate and the presence of a short accessory canal on the anterior dorsolateral plate. A new phylogenetic analysis positions Groenlandaspididae in a monophyly with the phlyctaeniid families Arctolepidae and Arctaspididae, however, the specific intrarelationships of groenlandaspidids remain poorly resolved., Competing Interests: The authors declare that they have no competing interests., (© 2024 Fitzpatrick et al.)

Yinlong downsi , the earliest known ceratopsian, is represented by dozens of specimens of different sizes collected from the Upper Jurassic of the Junggar Basin, northwestern China. Here, we present the first comprehensive study on the bone histology of Yinlong downsi based on ten specimens varying in size. Four ontogenetic stages are recognized: early juvenile, late juvenile, subadult, and adult. The reconstructed growth curve suggests that Yinlong may reach sexual maturity at 6 years old, which is earlier than that of the well-studied early-diverging ceratopsian Psittacosaurus (9 years old) but later than ceratopsids (about 3 to 5 years old). This may indicate that sexual maturity begins earlier during the evolution of ceratopsians, and that the giant size of ceratopsids is acquired by accelerating growth rates. The cortex of the tibia mainly consists of fibrolamellar bone tissues, but parallel-fibered bone and lines of arrested growth (LAGs) are very common throughout ontogeny, suggesting a moderate growth rate. Quantitative analysis indicates that Yinlong has a maximum growth rate similar to those of other small-sized dinosaurs such as Psittacosaurus , Dysalotosaurus , and Troodon , and their maximum growth rates are higher than those of extant squamates and crocodiles but lower than those of extant mammals and large dinosaurs. This suggests that body size plays a more important role in growth rate than other factors such as phylogenetic position and/or diet among non-avian dinosaurs., Competing Interests: The authors declare that they have no competing interests., (© 2024 Han et al.)

Pterosaurs were the first vertebrates to achieve powered flight. Early pterosaurs had long stiff tails with a mobile base that could shift their center of mass, potentially benefiting flight control. These tails ended in a tall, thin soft tissue vane that would compromise aerodynamic control and efficiency if it fluttered excessively during flight. Maintaining stiffness in the vane would have been crucial in early pterosaur flight, but how this was achieved has been unclear, especially since vanes were lost in later pterosaurs and are absent in birds and bats. Here, we use Laser-Stimulated Fluorescence imaging to reveal a cross-linking lattice within the tail vanes of early pterosaurs. The lattice supported a sophisticated dynamic tensioning system used to maintain vane stiffness, allowing the whole tail to augment flight control and the vane to function as a display structure., Competing Interests: NJ, TK, MH, TH, MP No competing interests declared, (© 2024, Jagielska et al.)

The Lower Jurassic Lufeng Formation of China has long been recognized for its diverse early-diverging sauropodomorph dinosaurs, with eight genera and ten species, representing more than half the Laurasian records. In this paper, we describe a new genus and species of non-sauropodan sauropodomorph, Lishulong wangi gen. et sp. nov., from Yunnan Province in southwestern China. This new taxon is represented by a partial skeleton including the skull and nine articulated cervical vertebrae, which differs from other Lufeng forms in both cranial and cervical characteristics. It bears several autapomorphies of the nasal process, the maxillary neurovascular foramen, and the cervical neural spine. Phylogenetic analysis reveals that Lishulong is an early-diverging member of the Sauropodiformes, and the sister-taxon of Yunnanosaurus . Elucidating the novel osteology of Lishulong , it possessed the largest sauropodomorph cranial material currently identified from the Lufeng Formation, not only enriches the diversity of the Lufeng dinosaur assemblage, but also enhances our understanding of the character evolution in early-diverging sauropodiforms. Furthermore, information about paleobiogeographic distributions indicates that Early Jurassic sauropodomorphs, especially Chinese taxa, have maintained multiple dispersions and exchanges within Pangaea., Competing Interests: The authors declare that they have no competing interests., (© 2024 Zhang et al.)

For over a century, there was a major gap in our understanding of the evolution of the flying Mesozoic reptiles, the pterosaurs, with a major morphological gap between the early forms and the derived pterodactyloids. 1 Recent discoveries have found a cluster of intermediate forms that have the head and neck of the pterodactyloids but the body of the early grade, 2 yet this still leaves fundamental gaps between these intermediates and both earlier and more derived pterosaurs. Here, we describe a new and large Jurassic pterosaur, Skiphosoura bavarica gen. et sp. nov., preserved in three dimensions, that helps bridge the gap between current intermediate pterosaurs and the pterodactyloids. A new phylogeny shows that there is a general progression of key characteristics of increasing head size, increasing length of neck and wing metacarpal, modification to the fifth toe that supports the rear wing membrane, and gradual reduction in tail length and complexity from earlier pterosaurs into the first pterodactyloids. This also shows a clear evolution of the increasing terrestrial competence of derived pterosaurs. Furthermore, this closes gaps between the intermediates and their ancestors and descendants, and it firmly marks the rhamphorhynchines and ctenochasmatid clades as, respectively, being the closest earliest and latest groups to this succession of transitional forms., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Recent years have seen increasing scientific interest in whether neuron counts can act as correlates of diverse biological phenomena. Lately, Herculano-Houzel (2023) argued that fossil endocasts and comparative neurological data from extant sauropsids allow to reconstruct telencephalic neuron counts in Mesozoic dinosaurs and pterosaurs, which might act as proxies for behaviors and life history traits in these animals. According to this analysis, large theropods such as Tyrannosaurus rex were long-lived, exceptionally intelligent animals equipped with "macaque- or baboon-like cognition", whereas sauropods and most ornithischian dinosaurs would have displayed significantly smaller brains and an ectothermic physiology. Besides challenging established views on Mesozoic dinosaur biology, these claims raise questions on whether neuron count estimates could benefit research on fossil animals in general. Here, we address these findings by revisiting Herculano-Houzel's (2023) work, identifying several crucial shortcomings regarding analysis and interpretation. We present revised estimates of encephalization and telencephalic neuron counts in dinosaurs, which we derive from phylogenetically informed modeling and an amended dataset of endocranial measurements. For large-bodied theropods in particular, we recover significantly lower neuron counts than previously proposed. Furthermore, we review the suitability of neurological variables such as neuron numbers and relative brain size to predict cognitive complexity, metabolic rate and life history traits in dinosaurs, coming to the conclusion that they are flawed proxies for these biological phenomena. Instead of relying on such neurological estimates when reconstructing Mesozoic dinosaur biology, we argue that integrative studies are needed to approach this complex subject., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Thescelosaurines are a group of early diverging, ornithischian dinosaurs notable for their conservative bauplans and mosaic of primitive features. Although abundant within the latest Cretaceous ecosystems of North America, their record is poor to absent in earlier assemblages, leaving a large gap in our understanding of their evolution, origins, and ecological roles. Here we report a new small bodied thescelosaurine-Fona herzogae gen. et sp. nov.-from the Mussentuchit Member of the Cedar Mountain Formation, Utah, USA. Fona herzogae is represented by multiple individuals, representing one of the most comprehensive skeletal assemblages of a small bodied, early diverging ornithischian described from North America to date. Phylogenetic analysis recovers Fona as the earliest member of Thescelosaurinae, minimally containing Oryctodromeus, and all three species of Thescelosaurus, revealing the clade was well-established in North America by as early as the Cenomanian, and distinct from, yet continental cohabitants with, their sister clade, Orodrominae. To date, orodromines and thescelosaurines have not been found together within a single North American ecosystem, suggesting different habitat preferences or competitive exclusion. Osteological observations reveal extensive intraspecific variation across cranial and postcranial elements, and a number of anatomical similarities with Oryctodromeus, suggesting a shared semi-fossorial lifestyle., (© 2024 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Much of our view on Mesozoic dinosaur diversity is obscured by biases in the fossil record. In particular, spatiotemporal sampling heterogeneity affects identification of the timing and geographical location of radiations, the recognition of the latitudinal diversity gradient, as well as interpretation of purported extinctions, faunal turnovers and their drivers, including the Early Jurassic Jenkyns Event and across the Jurassic/Cretaceous boundary. The current distribution of sampling means it is impossible to robustly determine whether these 'events' were globally synchronous and geologically instantaneous or spatiotemporally staggered. Accounting for sampling heterogeneity is also paramount to reconciling notable differences in results based on sampling-standardized dinosaur species richness versus reconstructions of diversification rates, particularly with regards to the lead-up to the Cretaceous/Paleogene mass extinction. Incorporation of a greater proportion of stratigraphically well-resolved dinosaurs into analyses is also imperative and must include the substantial Mesozoic radiation of birds. Given the relative rarity of temporally successive, well-sampled spatial windows, it remains possible that dinosaur species richness and diversification rate showed little change after the clade's initial radiation until the Cretaceous/Paleogene boundary. However, better understanding of underlying sampling, combined with a holistic approach to reconstructing dinosaur diversity and diversification, is an important step in testing this hypothesis.

A new specimen of Haplocanthosaurus is described based on bones excavated from the Late Jurassic Dry Mesa Dinosaur Quarry near Delta, Colorado. The specimen consists of seven dorsal vertebrae and a right tibia and is identified as Haplocanthosaurus based on the dorsally angled transverse processes, tall neural arch peduncles, low parapophyses relative to the diapophyses in the posterior dorsal vertebrae, and the robustness of the tibia combined with a greatly expanded distal articular surface. The discovery adds to our understanding of the biostratigraphy of Haplocanthosaurus, showing this genus is definitively present in the Brushy Basin Member of the Morrison Formation, and making this individual the geologically youngest Haplocanthosaurus specimen on the Colorado Plateau. The identification of this genus adds to the known diversity of sauropods at Dry Mesa Dinosaur Quarry (DMDQ), which is at least six distinct genera, making DMDQ the most diverse single locality of sauropods in the Morrison Formation and the world., (© 2024 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Our fascination with dinosaur brains and their capabilities essentially began with the first dinosaur discovery. The history of this study is a useful reflection of palaeoneurology as a whole and its relationship to a more inclusive evolutionary neuroscience. I argue that this relationship is imbued with high heuristic potential, but one whose realization requires overcoming certain constraints. These constraints include the need for a stable phylogenetic framework, methods for efficient and precise endocast construction, and fossil researchers who are steeped in a neuroscience perspective. The progress that has already been made in these areas sets the stage for a more mature palaeoneurology-not only one capable of being informed by neuroscience discoveries but one that drives such discoveries. I draw from work on the size, shape, behavioural correlates and developmental role of the dinosaur brain to outline current advances in dinosaur palaeoneurology. My examples largely are taken from theropods and centre on questions related to the origin of birds and their unique locomotory capabilities. The hope, however, is that these exemplify the potential for study in other dinosaur groups, and for utilizing the dinosaur-bird lineage as a parallel model on a par with mammals for studying encephalization.

The early radiation of dinosaurs remains a complex and poorly understood evolutionary event 1-4 . Here we use hundreds of fossils with direct evidence of feeding to compare trophic dynamics across five vertebrate assemblages that record this event in the Triassic-Jurassic succession of the Polish Basin (central Europe). Bromalites, fossil digestive products, increase in size and diversity across the interval, indicating the emergence of larger dinosaur faunas with new feeding patterns. Well-preserved food residues and bromalite-taxon associations enable broad inferences of trophic interactions. Our results, integrated with climate and plant data, indicate a stepwise increase of dinosaur diversity and ecospace occupancy in the area. This involved (1) a replacement of non-dinosaur guild members by opportunistic and omnivorous dinosaur precursors, followed by (2) the emergence of insect and fish-eating theropods and small omnivorous dinosaurs. Climate change in the latest Triassic 5-7 resulted in substantial vegetation changes that paved the way for ((3) and (4)) an expansion of herbivore ecospace and the replacement of pseudosuchian and therapsid herbivores by large sauropodomorphs and early ornithischians that ingested food of a broader range, even including burnt plants. Finally, (5) theropods rapidly evolved and developed enormous sizes in response to the appearance of the new herbivore guild. We suggest that the processes shown by the Polish data may explain global patterns, shedding new light on the environmentally governed emergence of dinosaur dominance and gigantism that endured until the end-Cretaceous mass extinction., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

The knowledge of Pliensbachian (Early Jurassic, ∼192.9-184.2 Ma) plesiosaurs is notoriously insufficient. Although there have been specimens described from different parts of the world, only three of them have been established as diagnosable taxa. Here, we describe two previously unreported lower Pliensbachian plesiosaur occurrences that originate from two sites located in North Rhine-Westphalia, Germany. One of the new occurrences is represented by three cervical and three indeterminable vertebrae from Werther, the other includes two associated pectoral or anterior dorsal vertebrae from Bielefeld. Although highly incomplete, the Werther individual, which derived from the Uptonia jamesoni Zone, is found to represent the only reliably identified early Pliensbachian pliosaurid known to date. Its material is geographically and stratigraphically proximate to the late Pliensbachian pliosaurid Arminisaurus schuberti , found in a clay-pit located in the Bielefeld district of Jöllenbeck. However, even though the Werther plesiosaur and A. schuberti show a broadly similar morphology of the preserved cervical section, a precise identification of the Werther taxon is currently impossible., Competing Interests: The authors declare there are no competing interests., (©2024 Sachs et al.)

Northeast China's Early Cretaceous Yixian Formation preserves spectacular fossils that have proved extraordinarily important in testing evolutionary hypotheses involving the origin of birds and the distribution of feathers among nonavian dinosaurs. These fossils occur either flattened with soft tissue preservation (including feathers and color) in laminated lacustrine strata or as three-dimensional (3D) skeletons in "life-like" postures in more massive deposits. The relationships of these deposits to each other, their absolute ages, and the origin of the extraordinary fossil preservation have been vigorously debated for nearly a half century, with the prevailing view being that preservation was linked to violent volcanic eruptions or lahars, similar to processes that preserved human remains at Pompeii. We present high-precision zircon U-Pb geochronology from cores and outcrops, demonstrating that Yixian Formation accumulation rates are more than an order of magnitude higher than usually estimated. Additionally, we provide zircon provenance and sedimentological data from 3D dinosaur fossils, which imply that their death and burial occurred in collapsed burrows, rather than via a catastrophic volcanogenic mechanism. In the studied area, the three principal fossil-rich intervals of the Yixian occur as a cyclic sequence that correspond to periods of high precipitation. Using Bayesian-Markov Chain Monte Carlo approaches, we constrain the total duration of the sequence to less than ~93,000 y and suggest that climatic precession paced the expression of these cyclic sediments. Rather than representing multiple, Pompeii-like catastrophes, the Yixian Formation is instead a brief snapshot of normal life and death in an Early Cretaceous continental community., Competing Interests: Competing interests statement:The authors declare no competing interest.

Avian evolution led to morphological adaptive variations in feet. Diverse foot types are accompanied by a diverse muscle system, allowing birds to adopt different primary lifestyles, and to display various locomotor and manipulative skills. We provide insights of evolutionary and functional significance on the avian foot architecture through Anatomical Network Analysis, a methodology focused on connectivity patterns of anatomical parts. Here, we show that: (1) anatomical parts largely conserved in living birds and already present in ancestral dinosaurs exhibit the highest connectivity degree, (2) there is no link between the more complex foot networks and the ability to perform more specialized skills or a higher number of different tasks, (3) there is a trend towards the simplification of foot networks on a macroevolutionary scale within birds, and (4) foot networks are phylogenetically constrained and conserved in all birds despite their foot type diversity, probably due to stabilizing selection., Competing Interests: Competing interests The authors declare no competing interests., (© 2024. The Author(s).)

Dinosaurs thrived for over 160 million years in Mesozoic ecosystems, displaying diverse ecological and evolutionary adaptations. Their ecology was shaped by large-scale climatic and biogeographic changes, calling for a 'deep-time' macroecological investigation. These factors include temperature fluctuations and the break up of Pangaea, influencing species richness, ecological diversity and biogeographic history. Recent improvements in the dinosaur fossil record have enabled large-scale studies of their responses to tectonic, geographic and climatic shifts. Trends in species diversity, body size and reproductive traits can now be analysed using quantitative approaches like phylogenetic comparative methods, machine learning and Bayesian inference. These patterns sometimes align with, but also deviate from, first-order macroecological rules (e.g. species-area relationship, latitudinal biodiversity gradient, Bergmann's rule). Accurate reconstructions of palaeobiodiversity and niche partitioning require ongoing taxonomic revisions and detailed anatomical descriptions. Interdisciplinary research combining sedimentology, geochemistry and palaeoclimatology helps uncover the environmental conditions driving dinosaur adaptations. Fieldwork in under-sampled regions, particularly at latitudinal extremes, is crucial for understanding the spatial heterogeneity of dinosaur ecosystems across the planet. Open science initiatives and online databases play a key role in advancing this field, enriching our understanding of deep-time ecological processes, and offering new insights into dinosaur macroecology and its broader implications.

Riojasuchus tenuisceps was a pseudosuchian archosaur from the Late Triassic period in Argentina. Like other ornithosuchids, it had unusual morphology such as a unique "crocodile-reversed" ankle joint, a lesser trochanter as in dinosaurs and a few other archosaurs, robust vertebrae, and somewhat shortened, gracile forelimbs. Such traits have fuelled controversies about its locomotor function-were its limbs erect or "semi-erect"? Was it quadrupedal or bipedal, or a mixture thereof? These controversies seem to persist because analyses have been qualitative (functional morphology) or correlative (morphometrics) rather than explicitly, quantitatively testing mechanistic hypotheses about locomotor function. Here, we develop a 3D whole-body model of R. tenuisceps with the musculoskeletal apparatus of the hindlimbs represented in detail using a new muscle reconstruction. We use this model to quantify the body dimensions and hindlimb muscle leverages of this enigmatic taxon, and to estimate joint ranges of motion and qualitative joint functions. Our model supports prior arguments that R. tenuisceps used an erect posture, parasagittal gait and plantigrade pes. However, some of our inferences illuminate the rather contradictory nature of evidence from the musculoskeletal system of R. tenuisceps-different features support (or are ambiguous regarding) quadrupedalism or bipedalism. Deeper analyses of our biomechanical model could move toward a consensus regarding ornithosuchid locomotion. Answering these questions would not only help understand the palaeobiology and bizarre morphology of this clade, but also more broadly if (or how) locomotor abilities played a role in the survival versus extinction of various archosaur lineages during the end-Triassic mass extinction event., (© 2024 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Archosauria originated around the Earth's largest biotic crisis that severely affected all ecosystems globally, the Permotriassic Mass extinction event, and comprises two crown-group lineages: the bird-lineage and the crocodylian lineage. The bird lineage includes the iconic pterosaurs, as well as dinosaurs and birds, whereas the crocodylian lineage includes clades such as aetosaurs, poposaurs, "rauisuchians," as well as Crocodylomorpha; the latter being represented today only by less than 30 extant species of Crocodylia. Despite playing important roles during Mesozoic and Cenozoic ecosystems, both on land and in water, Pseudosuchia received far less attention compared to the bird-lineage, which is also reflected in number and scope of histological studies so far. Lately, the field has seen a shift of focus toward pseudosuchians, however, and the symposium on "Paleohistological Inferences of Paleobiological Traits in Pseudosuchia" held during the International Congress of Vertebrate Morphology 2023 in Cairns, Queensland, Australia, is the latest proof of that. To put these novel aspects of paleohistological and paleobiological research into context, an overview of the non-extant pseudosuchian taxa whose postcranial bones were studied so far is provided here (c. 80 species out of a total of more than 700 extinct species described) and recent trends in pseudosuchian osteohistology are highlighted. In addition, histological studies on cranial and dental material and other potential hard tissues, such as eggshells and otoliths, are briefly reviewed as well., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Carcharodontosaurids were gigantic terrestrial dinosaurs and top predators of dinosaur faunas in Gondwanan landmasses during the "Mid"-Cretaceous Period. Despite their wide geographical and stratigraphical distribution, essential parts of their anatomy are still poorly known. The present contribution aims to describe a new partial skeleton of the carcharodontosaurid Taurovenator violantei, which was previously known only by an isolated postorbital bone coming from Cenomanian-Turonian beds of northern Patagonia, Argentina. The neck of Taurovenator is composed of notably high anterior cervicals, bearing neural spines with expanded, flange-like dorsal tips which are successively imbricated. This condition has been reported previously in the carcharodontosaurid Acrocanthosaurus, but its occurrence in Taurovenator and other members of the clade suggests it may represent a synapomorphy of this theropod family. This unique neck morphology was probably related to strong modifications in musculature and restriction in the range of movements within the neck, but not with the head. The new specimen also affords valuable anatomical information on the forelimb of Patagonian carcharodontosaurids. As in other giganotosaurines, Taurovenator shows strongly reduced forelimbs, particularly the forearm, showing hand elements with elongated non-ungual phalanges, and well-marked articular surfaces and muscular insertions, suggesting highly movable digits. This new specimen of Taurovenator allows us to expand anatomical and morpho-functional discussions about the carcharodontosaurid clade., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The Permian mesosaurs are well known for being the earliest amniotes to exhibit adaptations for living in a marine environment (Irati-Whitehill Sea). In addition to their set of skeletal features associated with aquatic dwelling life, their dentition includes important characteristics related to feeding in this habitat, which is described in this work, based on the analysis of mesosaur specimens from the Lower Permian Irati Formation of Brazil. Mesosaurs have several slender, conical teeth bordered by enamel apicobasal ridges, a feature predominantly found in aquatic amniotes. Internally, the dentine walls are formed by the arrangement of layers of orthodentine and globular dentine. To prevent tooth loss, the basal area is equipped with plicidentine, a particular type of orthodentine, allied with cementum, alveolar bone trabeculae, and periodontal space that reinforces anchorage and provides some flexibility. The teeth are replaced in a labio-vertical path, and the dentition replaces alternately. This feature is regarded as plesiomorphic, and it ensures the oral cavity is supplied with enough teeth. However, these features do not the assessment of whether mesosaurs teeth were capable of piercing prey with resistant tegument. Instead, we interpret this adaptation as a mechanism for catching prey, at least in adults, and we endorse a possible ontogenetic dietary shift from small to large forms., (© 2024 American Association for Anatomy.)

Dinosaurs potentially originated in the mid-palaeolatitudes of Gondwana 245-235 million years ago (Ma) and may have been restricted to cooler, humid areas by low-latitude arid zones until climatic amelioration made northern dispersals feasible ca 215 Ma. However, this scenario is challenged by new Carnian Laurasian fossils and evidence that even the earliest dinosaurs had adaptations for arid conditions. After becoming globally distributed in the Early-Middle Jurassic (200-160 Ma), dinosaurs experienced vicariance driven by Pangaean fragmentation. Regional extinctions and trans-oceanic dispersals also played a role, and the formation of ephemeral land connections meant that older vicariance patterns were repeatedly overprinted by younger ones, creating a reticulate biogeographic history. Palaeoclimates shaped dispersal barriers and corridors, including filters that had differential effects on different types of dinosaurs. Dinosaurian biogeographic research faces many challenges, not the least of which is the patchiness of the fossil record. However, new fossils, extensive databasing and improved analytical methods help distinguish signal from noise and generate fresh perspectives. In the future, developing techniques for quantifying and ameliorating sampling biases and modelling the dispersal capacities of dinosaurs are likely to be two of the key components in our modern research programme.

A new lithostrotian titanosaur, Qunkasaura pintiquiniestra gen. et sp. nov., is described based on a single partial skeleton from the late Campanian-early Maastrichtian fossil-site of Lo Hueco (Cuenca, Spain). This new taxon is supported by an exclusive combination of characters that highlights strong convergences with members of the South American Aeolosaurini. Qunkasaura allows to reorganise the complex phylogenetic relationships of the increasingly diverse finicretaceous sauropods of Europe. Phylogenetic analyses places Qunkasaura within Saltasauridae and possibly Opisthocoelicaudiinae, together with Abditosaurus. A new clade is established, Lohuecosauria, including Saltasaurus, Lohuecotitan, their most recent common ancestor and all its descendants. Two distinct Ibero-Armorican Campanian-Maastrichtian saltasauroid lineages are recognised: (i) Lirainosaurinae that is exclusive from Europe, and (ii) a saltasaurid lineage with possible opisthocoelicaudiine affinities, with a Laurasian distribution. Lirainosaurinae was a relict lineage including possible dwarf forms that evolved in isolation after reaching Europe before the Late Cretaceous through the Apulian route. The occurrence of opisthocoelicaudiines in Europe may be the result of a Late Cretaceous interchange between Europe and Asia. No evidence of insular dwarfism is found in the Ibero-Armorican opisthocoelicaudiines suggesting that they may have been newcomers to the area that arrived before the 'Maastrichtian Dinosaur Turnover' in southwestern Europe., (© 2024. The Author(s).)

Previous studies arguing for parental care in dinosaurs have been primarily based on fossil accumulations of adults and hatchlings, perinatal and post-hatchlings in nests and nest areas, and evidence of brooding, the majority of which date to the Late Cretaceous. Similarly, the general body proportions of preserved embryonic skeletons of the much older Early Jurassic Massospondylus have been used to suggest that hatchlings were unable to forage for themselves. Here, we approach the question of parental care in dinosaurs by using a combined morphological, chemical, and biomechanical approach to compare early embryonic and hatchling bones of the Early Jurassic sauropodomorph Lufengosaurus with those of extant avian taxa with known levels of parental care. We compare femora, the main weight-bearing limb bone, at various embryonic and post-embryonic stages in a precocious and an altricial extant avian dinosaur with those of embryonic and hatchling Lufengosaurus, and find that the rate and degree of bone development in Lufengosaurus is closer to that of the highly altricial Columba (pigeon) than the precocious Gallus (chicken), providing strong support for the hypothesis that Lufengosaurus was fully altricial. We suggest that the limb bones of Lufengosaurus hatchlings were not strong enough to forage for themselves and would likely need parental feeding., (© 2024. The Author(s).)

Derived ornithopods, such as hadrosaurids, show a high occurrence of fossilized lesions and diseases. However, paleopathologies in iguanodontians seem to be less common, considering the rich fossil record of these taxa in Europe, in particular in Belgium, Britain and Spain. Here, we describe an iguanodontian femur discovered in England that exhibits a large overgrowth of its lateral aspect, not previously recognized in any other similar remains. The specimen was scanned with micro-computed tomography (microCT) and later sectioned in three sites of the overgrowth for histological analysis. The femur belongs to an early adult Iguanodontia indet., based on the presence of a woven parallel fibered complex in the outer cortex and three to four lines of arrested growth. Internal analysis of the dome-like overgrowth suggests it can be diagnosed as a fracture callus. The injury might have negatively impacted upon the animal's locomotion as the trauma had occurred in the region above the knee, a crucial spot for hindlimb musculature. Finally, a cancellous medullary bone-like tissue was recognized in the medullary cavity next to the pathological overgrowth. An attempt was made to determine the precise nature of this tissue, as medullary bone is linked with the ovulation period in (avian) dinosaurs, whereas other types of endosteal, medullary bone-like tissue have previously been recognized in pathological bones., (© 2024 Anatomical Society.)

Background: Gastroliths are stones of uncertain purpose that are commonly found inside the rib cages of plesiosaur fossils worldwide. Gastroliths from four Alberta (Canada) plesiosaurs were studied to determine both their shapes and masses, and their mass fractions relative to body mass. One animal's set of gastroliths was 100% complete and fully visible, but the others showed varying degrees of loss, damage or obscuration, so estimations of their original states were needed., Methods: The studied animals were: Albertonectes vanderveldei , Fluvionectes sloanae , Nichollssaura borealis and Wapuskanectes betsynichollsae . The animals come from three different palaeoenvironments: open marine, near shore marine, and fluvial. Gastrolith shapes were classified as either xiphoid, cylindrical, discoidal or spherical based on observed and/or estimated dimensions. Although not all methods could be applied in all cases, gastrolith shapes and masses were estimated four different ways: (1) direct measurement and weighing of a subset and predicting the properties of the remaining obscured and hidden stones; (2) measuring triaxial ellipsoid dimensions of free stones to calculate volumes and multiplying by the mass density of chert; (3) measuring two visible triaxial dimensions of embedded stones, estimating the hidden third dimension three different ways, and then determining volumes and masses by calculation; and (4) predicting the density and mass of a densely packed cluster of small gastroliths using geometrical arguments., Results: Total gastrolith mass never exceeded 0.2% of body mass in any plesiosaur, and is consistent with the idea that the amounts of gastroliths recovered with plesiosaurs would be ineffective as ballast. The largest plesiosaur in the sample had the largest single gastrolith and total gastrolith mass increases with body size. The shape characteristics of the gastroliths were different for different environments, but compositionally they are dominated by black cherts. A possible common source for the gastroliths was identified for the two geographically close and near-contemporanous Nichollssaura and Wapuskanectes ., Competing Interests: The author declares that he has no competing interests., (© 2024 Henderson.)

Modern birds possess highly encephalized brains that evolved from non-avian dinosaurs. Evolutionary shifts in developmental timing, namely juvenilization of adult phenotypes, have been proposed as a driver of head evolution along the dinosaur-bird transition, including brain morphology. Testing this hypothesis requires a sufficient developmental sampling of brain morphology in non-avian dinosaurs. In this study, we harness brain endocasts of a postnatal growth series of the ornithischian dinosaur Psittacosaurus and several other immature and mature non-avian dinosaurs to investigate how evolutionary changes to brain development are implicated in the origin of the avian brain. Using three-dimensional characterization of neuroanatomical shape across archosaurian reptiles, we demonstrate that (i) the brain of non-avian dinosaurs underwent a distinct developmental trajectory compared to alligators and crown birds; (ii) ornithischian and non-avialan theropod dinosaurs shared a similar developmental trajectory, suggesting that their derived trajectory evolved in their common ancestor; and (iii) the evolutionary shift in developmental trajectories is partly consistent with paedomorphosis underlying overall brain shape evolution along the dinosaur-bird transition; however, the heterochronic signal is not uniform across time and neuroanatomical region suggesting a highly mosaic acquisition of the avian brain form., (© 2024. The Author(s).)

Ornithopod dinosaurs evolved numerous craniodental innovations related to herbivory. Nonetheless, the relationship between occlusion, tooth wear rate, and tooth replacement rate has been neglected. Here, we reconstruct tooth wear rates by measuring tooth replacement rates and tooth wear volumes, and document their dental microwear. We demonstrate that total tooth volume and rates of tooth wear increased steadily during ornithopod evolution, with deeply-nested taxa wearing up to 3360 mm 3 of tooth volume/day. Increased wear resulted in asymmetric tooth crown formation with uneven von Ebner line increment width by the Late Jurassic, and in faster tooth replacement rates in multiple lineages by the mid-Cretaceous. Microwear displays a contrasting pattern, with decreasing complexity and pit percentages in deeply-nested and later-occurring taxa. We hypothesize that early ornithopods were browsers and/or frugivores but deeply nested iguanodontians were bulk-feeders, eating tougher, less nutritious plants; these trends correlate with increasing body mass and longer gut passage times., (© 2024. The Author(s).)

The basal macronarian sauropod Europasaurus holgeri is known only from the Late Jurassic of the Langenberg Quarry near Goslar, Lower Saxony, Germany. Europasaurus has been identified as an insular dwarf and shows a clear resemblance to Camarasaurus and Giraffatitan . This study provides a detailed description of the dentition of Europasaurus based on an array of fossils outstanding in their abundance, variety of preservation, and ontogenetic range. Dental morphology for the replacement and functional dentitions, the tooth replacement pattern, and implications for food intake are described for the Europasaurus dentition, which is characterized by broad-crowned teeth. Characteristic features for Europasaurus are the presence of denticles on replacement teeth, the wrinkled enamel, and large wear facets both on the apex and on the carinae of the tooth crowns. The partially articulated skull SNHM-2207-R and isolated tooth rows DfmMh/FV 580.1 and DfmMh/FV 896.7 suggest the presence of strong connective tissue partially covering the teeth. This connective tissue would have provided stability and protection for the teeth. Evidence for this connective tissue include exposed tooth necks, in-situ teeth with strongly resorbed roots which no longer would have been connected to the jaw bone, and wrinkled enamel and its surface pattern. The same features can be observed in other sauropod taxa as well. We therefore suggest that eusauropods in general possessed this connective tissue structure, which may be an autapomorphy of the group. Possibly, this hypothetical structure is homologous to the rhamphotheca in birds and some non-avian theropods, which, however rarely, show such a close integration of keratinous tissue and teeth that we hypothesize here., Competing Interests: The authors declare that they have no competing interests., (© 2024 Régent et al.)

Therocephalia are an important clade of non-mammalian therapsids that evolved a diverse array of morphotypes and body sizes throughout their evolutionary history. The postcranial anatomy of therocephalians has largely been overlooked, but remains important towards understanding aspects of their palaeobiology and phylogenetic relationships. Here, we provide the first postcranial description of the large akidnognathid eutherocephalian Moschorhinus kitchingi by examining multiple specimens from fossil collections in South Africa. We also compare the postcranial anatomy with previously described therocephalian postcranial material and provide an updated literature review to ensure a reliable foundation of comparison for future descriptive work. Moschorhinus shares all the postcranial features of eutherocephalians that differentiate them from early-diverging therocephalians, but is differentiated from other eutherocephalian taxa by aspects concerning the scapula, interclavicle, sternum, manus, and femur. The novel anatomical data from this contribution shows that Moschorhinus possessed a stocky bauplan with a particularly robust scapula, humerus, and femur. These attributes, coupled with the short and robust skull bearing enlarged conical canines imply that Moschorhinus was well equipped to grapple with and subdue prey items. Additionally, the combination of these attributes differ from those of similarly sized coeval gorgonopsians, which would have occupied a similar niche in late Permian ecosystems. Moreover, Moschorhinus was the only large carnivore known to have survived the Permo-Triassic mass extinction. Thus, the subtle but important postcranial differences may suggest a type of niche partitioning in the predator guild during the Permo-Triassic mass extinction interval., Competing Interests: The authors declare that they have no competing interests., (© 2024 Stuart et al.)

The record of Mesozoic reptiles in the Northern region of Brazil is extremely limited, with the only definite occurrence consisting of two ziphodont teeth recovered from an oil well core in the municipality of Nova Olinda do Norte, Amazonas state, from strata of the Alter do Chão Formation (Cenomanian) of the Amazonas Basin. In this study, we aim to reevaluate MCT.R.514 using the most recent methodologies available for the identification of isolated theropod teeth. The cladistic analyses recovered MCT.R.514 as a possible metriacanthosaurid or non carcharodontosaurid allosauroid while the LDAs showed affinities between the specimen and Abelisauridae, Piatniktzysauridae, and Tyrannosauridae. MCT.R.514 was assigned as an abelisaurid due to its overall morphology that lacked metriacanthosaurid synapomorphies (i.e. spiraling mesial carina, strongly labially displaced distal carina, and well-marked mesiolingual longitudinal groove), while maintaining homoplastic features between both groups (i.e. irregular enamel texture, a lingually biconcave cross section, and a straight distal margin). This, together with a mostly Laurasian distribution of Metriacanthosauridae favors the assignment of MCT.R.514 as an abelisaurid as the most parsimonious hypothesis in this occasion. The identification of the specimen as an abelisaurid further expands the still scarce "Mid'' Cretaceous record of this clade in Brazil.

Take-off is a vital part of powered flight which likely constrains the size of birds, yet extinct pterosaurs are known to have reached far larger sizes. Three different hypothesised take-off motions (bipedal burst launching, bipedal countermotion launching, and quadrupedal launching) have been proposed as explanations for how pterosaurs became airborne and circumvented this proposed morphological limit. We have constructed a computational musculoskeletal model of a 5 m wingspan ornithocheiraean pterosaur, reconstructing thirty-four key muscles to estimate the muscle moment arms throughout the three hypothesised take-off motions. Range of motion constrained hypothetical kinematic sequences for bipedal and quadrupedal take-off motions were modelled after extant flying vertebrates. Across our simulations we did not find higher hindlimb moment arms for bipedal take-off motions or noticeably higher forelimb moment arms in the forelimb for quadrupedal take-off motions. Despite this, in all our models we found the muscles utilised in the quadrupedal take-off have the largest total launch applicable moment arms throughout the entire take-off sequences and for the take-off pose. This indicates the potential availability of higher leverage for a quadrupedal take-off than hypothesised bipedal motions in pterosaurs pending further examination of muscle forces., Competing Interests: Colin Palmer is an Academic Editor for PeerJ., (©2024 Griffin et al.)

Paleopathology, the study of diseases and injuries from the fossil record, allows for a unique view into the life of prehistoric animals. Pathologies have nowadays been described in nearly all groups of fossil vertebrates, especially dinosaurs. Despite the large number of skeletons, pathologies had never been reported in the sauropodomorph Plateosaurus trossingensis. Here we describe the first pathologies of Plateosaurus using two individuals with pathologies in the chevrons of the tail, from the Upper Triassic of Trossingen, SW Germany. The two specimens each contain three consecutive pathological chevrons. Our results show that the pathologies were caused by external trauma in one individual and potentially tendinous trauma in the other. Healing of the lesions allowed survival of both animals. Using additional pathological specimens found in other collections and from multiple localities, we observe that 14.8% of all individuals of Plateosaurus contain pathologies within their chevrons, suggesting it was a vulnerable bone., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Schaeffer et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Tyrannosaurids were the most derived group of Tyrannosauroidea and are characterized by having two body plans: gracile, long-snouted and robust, deep-snouted skulls. Both groups lived sympatrically in central Asia. Here, we report a new deep-snouted tyrannosaurid, Asiatyrannus xui gen. et sp. nov., from the Upper Cretaceous of Ganzhou City, southeastern China, which has produced the large-bodied and long-snouted Qianzhousaurus. Based on histological analysis, the holotype of Asiatyrannus xui is not a somatically mature adult, but it already passed through the most rapid growth stages. Asiatyrannus is a small to medium-sized tyrannosaurine, with a skull length of 47.5 cm and an estimated total body length of 3.5-4 m; or around half the size of Qianzhousaurus and other large-bodied tyrannosaurines in similar growth stages. Asiatyrannus and Qianzhousaurus are sympatric tyrannosaurid genera in the Maastrichtian of southeastern China. Asiatyrannus differs from Qianzhousaurus in that it has a proportionally deeper snout, longer premaxilla, deeper maxilla, and deeper dentary, and the cornual process of the lacrimal is inflated without developing a discrete horn. The different skull proportions and body sizes suggest that Asiatyrannus and Qianzhousaurus likely had different feeding strategies and occupied different ecological niches., (© 2024. The Author(s).)

A new troodontid dinosaur, Hypnovenator matsubaraetoheorum gen. et sp. nov., is described based on an articulated postcranial skeleton recovered from the fluvial deposits of the Albian Ohyamashimo Formation of the Sasayama Group in Tambasasayama City, Hyogo Prefecture, Japan. Hypnovenator is distinguished from other troodontids by four autapomorphies and a combination of additional features. Our phylogenetic analysis positions Hypnovenator as the oldest and one of the most basal troodontines, forming a clade with Gobivenator mongoliensis. The discovery of Hypnovenator suggests that small-bodied maniraptorans with a sleeping posture were common not only in environments with volcanic and eolian events or alluvial systems but also in fluvial systems. Geometric morphometric analysis of manual ungual phalanges shows that manual ungual phalanges I and III of Hypnovenator exhibit considerable morphological variation but are functionally similar, which differs from those of non-troodontine troodontids, reflecting the transition of manual motion within Troodontinae. Hypnovenator also has mosaic features in the pes related to cursoriality. This study reveals that asymmetrical arctometatarsus occurred by the Albian, and some morphological changes, such as shorter digit IV than digit III and non-ungual phalanges of digits III with roller joints and digit IV with weakly ginglymoid articulation, arose during the early Late Cretaceous., (© 2024. The Author(s).)

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., Competing Interests: David WE Hone is an Academic Editor for PeerJ, (© 2024 Habib and Hone.)

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., (© 2024. The Author(s).)

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., (© 2024. The Author(s).)

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., (© 2024. The Author(s).)

The Karoo Basin of South Africa is renowned for its abundance and diversity of therapsid fossils. Among the most ubiquitous and persistent of the Permian fauna is the small herbivorous dicynodont Diictodon feliceps . Intraspecific variation in Diictodon is historically confounding, and while ontogeny is frequently cited as a potential source of variation, observable developmental changes have never been calibrated. The present study revisits this issue, comparing three-dimensional landmark configurations of 82 Diictodon crania to investigate the association between shape, size and dimorphism. Beyond the statistically significant relationship between shape and allometry, our results determine the shape differences between juvenile and adult skulls of Diictodon , aligned with common craniofacial features documented in other tetrapod taxa. Functionally, these changes are attributed to development of the jaw musculature for feeding on larger, tougher plant matter during later ontogeny. Cranial morphological variation owing to sexual dimorphism is negligible, but distinct differences are noted in the allometric trajectories of each morphotype. A component of non-allometric variation cannot be accounted for, and we propose that this represents natural variation, rather than an artefact of taphonomic deformation.

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., Competing Interests: The authors declare there are no competing interests., (©2024 Hurtado et al.)