Basal Ornithischians Research Articles

Investigating the quadrupedal abilities of Scutellosaurus lawleri and its implications for locomotor behavior evolution among dinosaurs.

A reversion to secondary quadrupedality is exceptionally rare in nature, yet the convergent re-evolution of this locomotor style occurred at least four separate times within Dinosauria. Facultative quadrupedality, an intermediate state between obligate bipedality and obligate quadrupedality, may have been an important transitional step in this locomotor shift, and is proposed for a range of basal ornithischians and sauropodomorphs. Advances in virtual biomechanical modeling and simulation have allowed for the investigation of limb anatomy and function in a range of extinct dinosaurian species, yet this technique has not been widely applied to explore facultatively quadrupedal gait generation. This study places its focus on Scutellosaurus, a basal thyreophoran that has previously been described as both an obligate biped and a facultative quadruped. The functional anatomy of the musculoskeletal system (myology, mass properties, and joint ranges of motion) has been reconstructed using extant phylogenetic bracketing and comparative anatomical datasets. This information was used to create a multi-body dynamic locomotor simulation that demonstrates that whil quadrupedal gaits were physically possible, they did not outperform bipedal gaits is any tested metric. Scutellosaurus cannot therefore be described as an obligate biped, but we would predict its use of quadrupedality would be very rare, and perhaps restricted to specific activities such as foraging. This finding suggests that basal thyreophorans are still overwhelmingly bipedal but is perhaps indicative of an adaptive pathway for later evolution of quadrupedality.

Tooth replacement in Manidens condorensis: baseline study to address the replacement pattern in dentitions of early ornithischians

AbstractDental replacement in Heterodontosauridae has been debated over the last five decades primarily on indirect evidence, such as the development of wear facets and the position of erupted teeth. Direct observation of unerupted teeth provides unambiguous data for understanding tooth replacement but this has been done only for Heterodontosaurus and Fruitadens. This study addresses dental replacement in Manidens condorensis based on the positioning of functional and replacement teeth using microcomputed tomography data, differential wear along the dentition and the differences in labiolingual/apicobasal level of functional teeth. Dental replacement in Manidens condorensis was continuous in an anterior‐to‐posterior wave pattern, with asynchronous tooth eruption and the addition of new teeth posteriorly to the toothrow during ontogeny. Manidens shows the first evidence of dental replacement for the large dentary caniniform in Heterodontosauridae, which possibly had replacement timing distinct from the cheek dentition. Newly erupted teeth imbricate in a mesial cavity–distal crown base relationship during eruption, so that imbrication of the midposterior dentition remains unaltered during tooth replacement. The presence/absence of a small caniniform tooth in the D3 position of several specimens suggests possible intraspecific dimorphism in Manidens. On longitudinal sections of isolated crowns the histological features such as Howship's lacunae and odontoclast spaces are similar in size to extant reptiles. The differential wear decreasing posteriorly and hypothetical Z‐spacing below 2.3 in Manidens are similar to basal ornithischians. Tooth replacement in Heterodontosauridae (and other early ornithischians) provides key information for understanding the dynamics of jaw function and craniomandibular specialization to herbivory.

A new phylogeny of cerapodan dinosaurs

ABSTRACT This work attempts at providing a revised framework for ornithischian phylogeny, based on an exhaustive data compilation of already published analyses, a critical re-evaluation of osteological characters and an in-depth checking of characters scoring to fix mistakes that have accumulated in previous analyses; we have also included recently described basal ornithischians, marginocephalians and ornithopods. ‘Heterodontosaurids’ are recovered as a paraphyletic group of basal Marginocephalia that progressively lead to the dome-headed ‘true’ pachycephalosaurs. ‘Heterodontosaurids’ consequently fall within Pachycephalosauria sensu Sereno, 1998. The reconfiguration of basal cerapodan relationships pulls the origins of ornithopods to the earliest stages of the Jurassic. Based on the present analysis, we also discuss ornithopod relationships, with a particular focus on basal Iguanodontia. Tenontosaurus is found as the basalmost iguanodontian. The monophyly of Rhabdodontomorpha in a position more derived than Tenontosaurus is supported by the present analysis.

Cranial Musculature in Herbivorous Dinosaurs: A Survey of Reconstructed Anatomical Diversity and Feeding Mechanisms.

Herbivorous dinosaurs exhibited diverse cranial feeding mechanisms. Although osteological, microwear, and biomechanical research has revealed some of this diversity, the evolutionary reorientation of cranial musculature throughout nonavian herbivorous Dinosauria and its influence on feeding mechanisms requires more study. Here, cranial muscle reconstructions in herbivorous dinosaurs are reviewed and informative anatomical characters are compared across 142 dinosaur genera (84 ornithischians, 36 sauropodomorphs, and 22 herbivorous nonavian theropods), both through examination of specimens and literature. Traits include those relating to the temporal region, adductor chamber, palate, and mandibular attachments, such as the coronoid elevation and retroarticular process. Findings reveal many combinations of anatomical traits influencing a diversity of feeding mechanisms. Some primarily more orthal feeders, including herbivorous theropods, nonsauropod sauropodomorphs, basal ornithischians, and derived stegosaurs (which also show varying degrees of coinciding slight palinal motion and long-axis hemimandibular rotation), possess traits indicative of more prominent temporal musculature and moderately sized palatal musculature. However, orthal feeding sauropods and pachycephalosaurs possess traits indicative of greatly reduced, low-angled temporal musculature, and enhanced palatal musculature producing a primarily vertical, orthal feeding vector. Among ankylosaurs, hadrosaurids, and neoceratopsians, a rostrolabial temporal muscle expansion is present (with a tall coronoid elevation in hadrosaurids and ceratopsids) for greater temporal muscle support and mechanical advantage for complex palinal feeding motions. This also aids in long-axis hemimandibular rotation against the predentary in hadrosaurs and ankylosaurs. This diversity in cranial muscle architecture provides an informative spectrum of numerous adaptations acquired given the evolution of various anatomical constraints in the skull. Anat Rec, 303:1104-1145, 2020. © 2019 American Association for Anatomy.

New data on the microvertebrate fauna from the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual Syncline, eastern Morocco)

AbstractThe Middle Jurassic – Early Cretaceous period witnessed the emergence of some major representatives of modern continental vertebrate groups (stem lissamphibians, squamates, therian mammals and birds) and angiosperms, at a time when fragmentation of Pangaea was underway. The successive Moroccan microvertebrate faunas of Ksar Metlili (?Berriasian) and Guelb el Ahmar (Bathonian) from the Anoual Syncline significantly improve our poor knowledge of Gondwanan and especially African palaeobiodiversity at this time. They are among the richest known from the Mesozoic of Gondwana, and are well placed in northwestern Africa to record faunal interchanges with Laurasia. Here we focus on the Ksar Metlili fauna, first documented in the 1980s and most recently resampled in 2010, which produced 24 541 microremains representing 47 species of 8 main groups (Chondrichthyes, Actinopterygii, Sarcopterygii, Lissamphibia, Lepidosauromorpha, Testudinata, Archosauromorpha and Synapsida). It includes remarkable taxa: the oldest stem boreosphenidan mammals from Gondwana, probably some of the last non-mammaliaform cynodonts, a basal ornithischian, possibly freshwater teleosaurid crocodylomorphs, and some of the rare occurrences of choristoderes and albanerpetontids in Gondwana. Comparison of the Ksar Metlili fauna with that of Guimarota (Kimmeridgian, Portugal) further provides evidence of numerous shared taxa of Laurasian affinities, in contrast to the occurrence of few taxa with Gondwanan affinities. This suggests complex palaeobiogeographical relationships – implying both vicariance and dispersal events – of North Africa within Gondwana at the Jurassic–Cretaceous transition. Finally, the faunal similarities with the Guelb el Ahmar fauna question the Cretaceous age of the Ksar Metlili fauna, suggesting an alternative possible Late Jurassic age.

Variability in the Evolution of Ornithischian Cranial Morphology

Phylogenetic relationships within Dinosauria have been recently called into question, including shifts among major taxonomic groups such as Ornithischia. As phylogenetic hypotheses change, so do hypotheses of morphological character evolution. Ornithischian dinosaurs exhibit a wide variety of cranial morphologies. They range from generalized ancestral morphologies, to derived and varied skull shapes with frills, thickened domes, or complex dental batteries associated with complex jaw joints. Here, I describe the evolution of cranial morphology within Ornithischia using an updated phylogenetic analysis and ancestral character state reconstruction methods to map changes in cranial characters through time. With an updated phylogenetic analysis of Ornithischia, I used both parsimony and maximum likelihood methods for ancestral state reconstructions to determine the changes in cranial characters. The ancestral reconstructions also allowed for a broader view of cranial morphological change through time. These estimates demonstrated that basal ornithischians, including those at the base of major clades (e.g. Thyreophora, Ornithopoda), maintained skull morphologies not unlike other basal dinosaurs outside of Ornithischia. The thyreophoran and neornithischian skulls show similar changes in cranial character evolution: relative reduction of the quadratojugal, general lengthening of the preorbital skull, including modifications to the premaxilla and nasal, an initial reduction in the size, placement, and shape of the antorbital fenestra, and incorporation of the palpebral in the supraorbital margin. While slight changes in the postorbital skull of thyreophorans were noted, the morphology within Neornithischia was markedly different. In addition to well‐known modifications of the parietal and squamosal in marginocephalians, other noted changes include movement of cranial foramina in the occipital and basal regions of the skull. This is in contrast to most ornithopods, whose skull bases and occipital regions vary relatively little from basal neornithischians. Derived ornithopods (e.g., hadrosaurs) demonstrate elaborate preorbital modifications, particularly involving the premaxilla and nasal. The dentition and mandible demonstrate modifications throughout the ornithischian tree. However, dramatic changes are noted among derived ornithopods with the development of massive dental batteries and laterally offset, handle‐shaped coronoid processes. This occurs to a lesser degree in ceratopsians as well. These changes in cranial morphology, while present in all ornithischian clades, varies in terms of rates of change. Within Thyreophora, a group that persisted for nearly 150 million years, the general changes within the skull and mandible are relatively small when compared to ornithopods. Ceratopsians and ornithopods demonstrate the most dramatic changes in overall cranial morphology, but ornithopods show the most remarkable alterations in mandibular anatomy through time. With these results, the analysis presented here may be used to develop new hypotheses for the selective pressures behind the varied evolution of the ornithischian skull.Support or Funding InformationUniversity of Iowa Max and Lorraine Littlefield Fund and T. Anne Cleary International Dissertation Research FellowshipThis abstract is from the Experimental Biology 2019 Meeting. There is no full text article associated with this abstract published in The FASEB Journal.

The Anatomical and Functional Evolution of the Femoral Fourth Trochanter in Ornithischian Dinosaurs.

The femoral fourth trochanter is the attachment site of the caudofemoralis musculature, which is the primary hindlimb retractor in most non-avian dinosaurs. Early ornithischian dinosaurs are uniquely characterized by a fourth trochanter with a prominent pendant process. Throughout the evolution of ornithischians, the fourth trochanter repeatedly converged on two major morphological changes: (1) the distal migration of the trochanter down the femoral shaft and (2) the loss of the pendant process. Both changes, as well as the original evolution of the pendant form, relate to a single major functional shift emphasizing caudofemoral leverage. Direct evidence of muscle scarring across the surface of the pendant process affirms that it served to extend the attachment of the primary caudofemoralis brevis tendon distally. A proximally located fourth trochanter is the basal condition in dinosaurs and other archosaurs, and the development of a pendant process lengthened the functional lever arm with regard to the insertion of the caudofemoralis. This adaptation afforded improved mechanical advantage, perhaps beneficial in the context of the newly assumed herbivorous diets of basal ornithischians. As some derived ornithischians increased in body size, a high-leverage system with a more distal caudofemoralis attachment evolved. In some groups, the fourth trochanter as a whole descended down the femur, eventually reaching a point where the pendant process was unnecessary. Sauropodomorphs, the other great lineage of dinosaur herbivores, converged on the same high-leverage distal fourth trochanter arrangement, but without first transitioning through a prominent pendant form. Anat Rec, 303:1146-1157, 2020. © 2019 Wiley Periodicals, Inc.

On the findings of dinosaurs in the Crimea

Ornithopod dinosaur Riabininohadros weberae from the Upper Cretaceous (upper Maastrichtian) of the Crimean Peninsula (Besh-Kosh) shows a set of primitive characters of hind limbs observed in iguanodontids and basal ornithischians and is referred to as Styracosterna indet. The second dinosaur specimen from Crimea (Aleshino) is a fragmentary skeleton, including cervical and dorsal vertebrae. It possibly belongs to advanced iguanodontids or primitive hadrosauroids. Thus, in the Maastrichtian of the Crimean Peninsula, at least two dinosaur species coexisted.

New Reconstruction of Cranial Musculature in Ornithischian Dinosaurs: Implications for Feeding Mechanisms and Buccal Anatomy.

The charismatic and diverse ornithischian dinosaurs exhibited some of the most extreme examples of cranial anatomy, inspiring decades of investigation into their muscular anatomy. Current ornithischian jaw muscle reconstructions, although parsimonious, pose concerns of small adductor muscles and caudally displaced insertions relative to mandibular proportions. Here, craniomandibular material of ornithischian genera spanning all subclades is reexamined for osteological correlates indicative of intracranial and oral soft tissues. M. adductor mandibulae externus (mAME) has traditionally been reconstructed as solely inserting along the caudal margin of the coronoid process for jaw closure. Here, a new mAME reconstruction is proposed in derived ornithischians, with the superficial-most mAME layer reconstructed as a rostrolabial expansion of muscle, exiting the cranium rostroventrally beneath a unique, laterally flaring jugal and inserting along the lateral surface of the coronoid process and its rostrally extending, shelf-like labial dentary ridge (LDR). Through previous dental microwear and morphological studies, ceratopsians, hadrosaurids, and ankylosaurs are known to have implemented a major palinal feeding component in their jaw motions, unlike other primarily basal ornithischians. This rostral fan-like extension of muscle in these derived clades would create a greater mandibular support system and mechanical advantage along the labial margin of the jaw, cradling the entire mandible while lifting it up into occlusion and retracting it. In hadrosaurids and ankylosaurs, this rostrolabially expanding muscle also acts in medial rotation of the dentaries about their long axes. With these new reconstructions, the notion of a novel, unparsimonious "cheek" muscle is rejected, with further discussion of plausible buccal soft tissues. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 303:347-362, 2020. © 2018 American Association for Anatomy.

Tooth development, histology, and enamel microstructure in Changchunsaurus parvus: Implications for dental evolution in ornithopod dinosaurs.

The great diversity of dinosaurian tooth shapes and sizes, and in particular, the amazing dental complexity in derived ornithischians has attracted a lot of attention. However, the evolution of dental batteries in hadrosaurids and ceratopsids is difficult to understand without a broader comparative framework. Here we describe tooth histology and development in the "middle" Cretaceous ornithischian dinosaur Changchunsaurus parvus, a small herbivore that has been characterized as an early ornithopod, or even as a more basal ornithischian. We use this taxon to show how a "typical" ornithischian dentition develops, copes with wear, and undergoes tooth replacement. Although in most respects the histological properties of their teeth are similar to those of other dinosaurs, we show that, as in other more derived ornithischians, in C. parvus the pulp chamber is not invaded fully by the newly developing replacement tooth until eruption is nearly complete. This allowed C. parvus to maintain an uninterrupted shearing surface along a single tooth row, while undergoing continuous tooth replacement. Our histological sections also show that the replacement foramina on the lingual surfaces of the jaws are likely the entry points for an externally placed dental lamina, a feature found in many other ornithischian dinosaurs. Surprisingly, our histological analysis also revealed the presence of wavy enamel, the phylogenetically earliest occurrence of this type of tissue. This contradicts previous interpretations that this peculiar type of enamel arose in association with more complex hadrosauroid dentitions. In view of its early appearance, we suggest that wavy enamel may have evolved in association with a shearing-type dentition in a roughly symmetrically-enameled crown, although its precise function still remains somewhat of a mystery.

New Data on Dinosaurs of the Crimean Peninsula.

Reexamination of the holotype of Riabininohadros weberae from the Upper Cretaceous (upper Maastrichtian) of the Crimean Peninsula (Besh-Kosh) allowed determination of previously unknown elements of the femur, astragalus, and calcaneus. This taxon shows a set of primitive characters observed in iguanodontids and basal ornithischians and is referred to as Styracosterna indet. The second dinosaur specimen from Crimea (Aleshino) is a fragmentary skeleton, including cervical and dorsal vertebrae. It possibly belongs to advanced iguanodontids or primitive hadrosauroids. Thus, in the Maastrichtian of the Crimean Peninsula, at least two dinosaur species coexisted.

Postcranial anatomy of Yinlong downsi (Dinosauria: Ceratopsia) from the Upper Jurassic Shishugou Formation of China and the phylogeny of basal ornithischians

Ceratopsia includes some of the best-known ornithischian dinosaurs. Many species are erected based on cranial elements alone, and the postcranial skeletons are either missing or undescribed in many taxa. Here we provide the first detailed postcranial description of Yinlong downsi based on the holotype and eight other well-preserved skeletons. Yinlong downsi from the early Late Jurassic Shishugou Formation of the Wucaiwan area, Xinjiang, China, represents one of the most basal ceratopsians. The detailed study of the postcranial skeleton reveals one feature unique to it among ceratopsians: a blade-like prepubic process of the pubis with an elongate notch near its ventral margin. The postcranial material of Yinlong shares some unique features with that of the ornithischian Stenopelix valdensis from the Early Cretaceous of Germany, and provides further evidence that the latter is a basal ceratopsian. A comprehensive phylogenetic analysis of basal ornithischians was built based on 72 taxa and 380 characters. Most of the characters are illustrated for the first time in order to clarify character states. The new ornithischian phylogeny confirms that Yinlong belongs to Chaoyangsauridae. Chaoyangsaurids and Psittacosaurus form a monophyletic group that is sister to all other ceratopsians. The new phylogeny also supports Stenopelix valdensis as a basal ceratopsian, and Mosaiceratops to be close to Coronosauria. Additionally, the new phylogeny agrees with other recent analyses that place heterodontosaurids as the most basal ornithischians rather than with marginocephalians. Furthermore, Isaberrysaura, which has been hypothesized to be a basal ornithopod, is recovered as one of the most basal stegosaurs for the first time. The former ‘hypsilophodontid’ taxa are recovered within Ornithopoda rather than outside Cerapoda, and Jeholosauridae is shown to be valid in this analysis.

Phylogenetic reassessment of Pisanosaurus mertii Casamiquela, 1967, a basal dinosauriform from the Late Triassic of Argentina

Pisanosaurus mertii was originally described on the basis of an incomplete skeleton from the early Late Triassic (Carnian) of northern Argentina. It is consistently regarded by most authors as a very basal ornithischian, the sister group of remaining members of the clade. The referral to Ornithischia is based mainly on tooth-bearing bones and tooth morphology. On the other hand, the postcranium is recognized as strikingly plesiomorphic for ornithischians, and even for dinosaurs. The recent description of non-dinosaurian dinosauriforms of the clade Silesauridae having ornithischian-like dentition invites a review of the phylogenetic affinities of Pisanosaurus. In this regard, an overview of the holotype specimen allows a reanalysis of previous anatomical interpretations of this taxon. The phylogenetic analysis presented here suggests that Pisanosaurus may be better interpreted as a member of the non-dinosaurian Silesauridae. It shares with silesaurids reduced denticles on the teeth, teeth fused to maxilla and dentary bone, sacral ribs shared between two sacral vertebrae, lateral side of proximal tibia with a fibular flange, and dorsoventrally flattened pedal ungual phalanges. The present analysis indicates that Pisanosaurus should be removed from the base of the Ornithischia and should no longer be considered the oldest representative of this dinosaurian clade.

Morphometric analysis of the forelimb and pectoral girdle of the Cretaceous ornithopod dinosaur Oryctodromeus cubicularis and implications for digging

ABSTRACTThe basal ornithopod Oryctodromeus cubicularis was described as burrowing due to its discovery in a burrow structure, and the presence of several morphological features considered consistent with burrowing. Using traditional and geometric morphometric analyses, the morphology of the humerus and scapula of ornithopods, basal ornithischians, and marginocephalians was analyzed to describe and characterize quantitatively the differences between Oryctodromeus and other ornithopods. These differences were then compared with the morphological adaptations for digging in mammals, because there are no adequate analogues for burrowing in extant archosaurs. A canonical variates analysis was also conducted on the geometric morphometric data to determine if phylogeny impacted morphological trends. The humerus of Oryctodromeus is slightly more robust than other basal ornithopods, indicating an adaptation for increased force applied to the humerus. The scapula provides the most compelling morphological support for digging in Oryctodromeus. The large acromion process and prominent scapular spine of Oryctodromeus, as well as the large posteroventrally expanded scapular blade, distinguish the scapula of Oryctodromeus from those of other ornithopods and would have provided surface areas for the attachment of the supracoracoideus, deltoideus clavicularis, and deltoideus scapularis muscles, the latter of which is important in digging in mammals. These features provide evidence for specialization for producing burrows by scratch digging.SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Complete forelimb myology of the basal theropod dinosaur Tawa hallae based on a novel robust muscle reconstruction method.

The forelimbs of nonavian theropod dinosaurs have been the subject of considerable study and speculation due to their varied morphology and role in the evolution of flight. Although many studies on the functional morphology of a limb require an understanding of its musculature, comparatively little is known about the forelimb myology of theropods and other bipedal dinosaurs. Previous phylogenetically based myological reconstructions have been limited to the shoulder, restricting their utility in analyses of whole-limb function. The antebrachial and manual musculature in particular have remained largely unstudied due to uncertain muscular homologies in archosaurs. Through analysis of the musculature of extant taxa in a robust statistical framework, this study presents new hypotheses of homology for the distal limb musculature of archosaurs and provides the first complete reconstruction of dinosaurian forelimb musculature, including the antebrachial and intrinsic manual muscles. Data on the forelimb myology of a broad sample of extant birds, crocodylians, lizards, and turtles were analyzed using maximum likelihood ancestral state reconstruction and examined together with the osteology of the early theropod Tawa hallae from the Late Triassic of New Mexico to formulate a complete plesiomorphic myology for the theropod forelimb. Comparisons with previous reconstructions show that the shoulder musculature of basal theropods is more similar to that of basal ornithischians and sauropodomorphs than to that of dromaeosaurids. Greater development of the supracoracoideus and deltoideus musculature in theropods over other bipedal dinosaurs correlates with stronger movements of the forelimb at the shoulder and an emphasis on apprehension of relatively large prey. This emphasis is further supported by the morphology of the antebrachium and the intrinsic manual musculature, which exhibit a high degree of excursion and a robust morphology well-suited for powerful digital flexion. The forelimb myology of Tawa established here helps infer the ancestral conformation of the forelimb musculature and the osteological correlates of major muscle groups in early theropods. These data are critical for investigations addressing questions relating to the evolution of specialized forelimb function across Theropoda.

Jaw muscle reconstruction and vector analysis in ornithischian dinosaurs (17.4)

Jaw muscle reconstruction has long been utilized for inferences in the paleobiology of ornithischian dinosaur feeding mechanisms. Previous studies in dinosaur jaw musculature have used extant phylogenetic bracketing methods to qualitatively compare muscle origins and insertions in a select few ornithischian genera. The current study uses criteria from previous jaw muscle studies, as well as more in‐depth case‐by‐case analyses, to reconstruct musculature and measure vector angles of the m. adductor mandibulae externus (MAME) complex as well as m. adductor mandibulae posterior (MAMP) in a large diversity of ornithischian taxa spanning all subclades. Results of both MAME and MAMP vectors show general trends from a mid‐range caudodorsal orientation in basal ornithischians and most armored thyreophorans to a decrease in vector angles indicating more caudally oriented jaw movements convergently in other taxa (e.g., derived thyreophorans, basal ornithopods, lambeosaurines, pachycephalosaurs, and derived ceratopsids). The extensive amount of variation in muscle vectors, both between and among subclades, however, suggests highly variable feeding mechanisms among genera and shows that feeding mechanisms of an entire subclade should not be based only on a single taxon, as has been previously done.

Postcranial anatomy of Jeholosaurus shangyuanensis (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China

ABSTRACT Jeholosaurus shangyuanensis is a small ornithischian dinosaur from the Lower Cretaceous Yixian Formation of the Lujiatun locality, Liaoning Province, China. Here, we provide the first detailed description of its postcranial skeleton based on the holotype and four other well-preserved skeletons, and compare it with material of other primitive cerapodans. Jeholosaurus can be diagnosed on the basis of one postcranial autapomorphy, relating to the absence of parapophyses from the anterior dorsal vertebrae, and a unique combination of postcranial characters, but its anatomy is otherwise similar to that of many other basal ornithischians. A phylogenetic analysis incorporating these new postcranial data confirms previous suggestions that Jeholosaurus is a basal ornithopod and that it forms a clade with Changchunsaurus and Haya; Koreanosaurus and Yueosaurus might also belong to this clade, though additional material of both will be required to test this hypothesis. The name Jeholosauridae is proposed for this apparently endemic group of Cretaceous East Asian taxa.

Computational modelling of locomotor muscle moment arms in the basal dinosaur Lesothosaurus diagnosticus: assessing convergence between birds and basal ornithischians

Ornithischia (the 'bird-hipped' dinosaurs) encompasses bipedal, facultative quadrupedal and quadrupedal taxa. Primitive ornithischians were small bipeds, but large body size and obligate quadrupedality evolved independently in all major ornithischian lineages. Numerous pelvic and hind limb features distinguish ornithischians from the majority of other non-avian dinosaurs. However, some of these features, notably a retroverted pubis and elongate iliac preacetabular process, appeared convergently in maniraptoran theropods, and were inherited by their avian descendants. During maniraptoran/avian evolution these pelvic modifications led to significant changes in the functions of associated muscles, involving alterations to the moment arms and the activation patterns of pelvic musculature. However, the functions of these features in ornithischians and their influence on locomotion have not been tested and remain poorly understood. Here, we provide quantitative tests of bipedal ornithischian muscle function using computational modelling to estimate 3D hind limb moment arms for the most complete basal ornithischian, Lesothosaurus diagnosticus. This approach enables sensitivity analyses to be carried out to explore the effects of uncertainties in muscle reconstructions of extinct taxa, and allows direct comparisons to be made with similarly constructed models of other bipedal dinosaurs. This analysis supports some previously proposed qualitative inferences of muscle function in basal ornithischians. However, more importantly, this work highlights ambiguities in the roles of certain muscles, notably those inserting close to the hip joint. Comparative analysis reveals that moment arm polarities and magnitudes in Lesothosaurus, basal tetanuran theropods and the extant ostrich are generally similar. However, several key differences are identified, most significantly in comparisons between the moment arms of muscles associated with convergent osteological features in ornithischians and birds. Craniad migration of the iliofemoralis group muscles in birds correlates with increased leverage and use of medial femoral rotation to counter stance phase adduction moments at the hip. In Lesothosaurus the iliofemoralis group maintains significantly higher moment arms for abduction, consistent with the hip abduction mode of lateral limb support hypothesized for basal dinosaurs. Sensitivity analysis highlights ambiguity in the role of musculature associated with the retroverted pubis (puboischiofemoralis externus group) in ornithischians. However, it seems likely that this musculature may have predominantly functioned similarly to homologous muscles in extant birds, activating during the swing phase to adduct the lower limb through lateral rotation of the femur. Overall the results suggest that locomotor muscle leverage in Lesothosaurus (and by inference basal ornithischians in general) was more similar to that of other non-avian dinosaurs than the ostrich, representing what was probably the basal dinosaur condition. This work thereby contradicts previous hypotheses of ornithischian-bird functional convergence.

Osteological correlates for quadrupedality in ornithischian dinosaurs

The evolution of quadrupedality from bipedal ancestors is an exceptionally rare transition in tetrapod evolution, but it has occurred several times within the herbivorous dinosaur clade Ornithischia. Stegosauria, Ankylosauria, and Ceratopsidae are all uncontroversially quadrupedal, while basal ornithischians and basal ornithopods are uncontroversially bipedal. However, stance in iguanodontian ornithopods, including the hadrosaurs, and in non-ceratopsid ceratopsians is debated because robust osteological correlates of quadrupedality have not been identified. We examine a suite of characteristics that have been previously proposed as osteological correlates for bipedality or quadrupedality in dinosaurs. These include both discrete anatomical features, which we assess as correlates for quadrupedality using character optimization onto a composite cladogram, and proportional ratios, which we assess as correlates by reconstructing nodal ancestral states using squared-change parsimony, followed by optimization. ...

The origin and early evolution of dinosaurs

The oldest unequivocal records of Dinosauria were unearthed from Late Triassic rocks (approximately 230 Ma) accumulated over extensional rift basins in southwestern Pangea. The better known of these are Herrerasaurus ischigualastensis, Pisanosaurus mertii, Eoraptor lunensis, and Panphagia protos from the Ischigualasto Formation, Argentina, and Staurikosaurus pricei and Saturnalia tupiniquim from the Santa Maria Formation, Brazil. No uncontroversial dinosaur body fossils are known from older strata, but the Middle Triassic origin of the lineage may be inferred from both the footprint record and its sister-group relation to Ladinian basal dinosauromorphs. These include the typical Marasuchus lilloensis, more basal forms such as Lagerpeton and Dromomeron, as well as silesaurids: a possibly monophyletic group composed of Mid-Late Triassic forms that may represent immediate sister taxa to dinosaurs. The first phylogenetic definition to fit the current understanding of Dinosauria as a node-based taxon solely composed of mutually exclusive Saurischia and Ornithischia was given as "all descendants of the most recent common ancestor of birds and Triceratops". Recent cladistic analyses of early dinosaurs agree that Pisanosaurus mertii is a basal ornithischian; that Herrerasaurus ischigualastensis and Staurikosaurus pricei belong in a monophyletic Herrerasauridae; that herrerasaurids, Eoraptor lunensis, and Guaibasaurus candelariensis are saurischians; that Saurischia includes two main groups, Sauropodomorpha and Theropoda; and that Saturnalia tupiniquim is a basal member of the sauropodomorph lineage. On the contrary, several aspects of basal dinosaur phylogeny remain controversial, including the position of herrerasaurids, E. lunensis, and G. candelariensis as basal theropods or basal saurischians, and the affinity and/or validity of more fragmentary taxa such as Agnosphitys cromhallensis, Alwalkeria maleriensis, Chindesaurus bryansmalli, Saltopus elginensis, and Spondylosoma absconditum. The identification of dinosaur apomorphies is jeopardized by the incompleteness of skeletal remains attributed to most basal dinosauromorphs, the skulls and forelimbs of which are particularly poorly known. Nonetheless, Dinosauria can be diagnosed by a suite of derived traits, most of which are related to the anatomy of the pelvic girdle and limb. Some of these are connected to the acquisition of a fully erect bipedal gait, which has been traditionally suggested to represent a key adaptation that allowed, or even promoted, dinosaur radiation during Late Triassic times. Yet, contrary to the classical "competitive" models, dinosaurs did not gradually replace other terrestrial tetrapods over the Late Triassic. In fact, the radiation of the group comprises at least three landmark moments, separated by controversial (Carnian-Norian, Triassic-Jurassic) extinction events. These are mainly characterized by early diversification in Carnian times, a Norian increase in diversity and (especially) abundance, and the occupation of new niches from the Early Jurassic onwards. Dinosaurs arose from fully bipedal ancestors, the diet of which may have been carnivorous or omnivorous. Whereas the oldest dinosaurs were geographically restricted to south Pangea, including rare ornithischians and more abundant basal members of the saurischian lineage, the group achieved a nearly global distribution by the latest Triassic, especially with the radiation of saurischian groups such as "prosauropods" and coelophysoids.