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The secondary evolution of quadrupedality from bipedal ancestry is a rare evolutionary transition in tetrapods yet occurred convergently at least three times within ornithischian dinosaurs. Despite convergently evolving quadrupedal gait, ornithischians exhibited variable anatomy, particularly in the forelimbs, which underwent a major functional change from assisting in foraging and feeding in bipeds to becoming principal weight-bearing components of the locomotor system in quadrupeds. Here, we use three-dimensional multi-body dynamics models to demonstrate quantitatively that different quadrupedal ornithischian clades evolved distinct forelimb musculature, particularly around the shoulder. We find that major differences in glenohumeral abduction–adduction and long axis rotation muscle leverages were key drivers of mechanical disparity, thereby refuting previous hypotheses about functional convergence in major clades. Elbow muscle leverages were also disparate across the major ornithischian lineages, although high elbow extension muscle leverages were convergent between most quadrupeds. Unlike in ornithischian hind limbs, where differences are more closely tied to functional similarity than phylogenetic relatedness, mechanical disparity in ornithischian forelimbs appears to have been shaped primarily by phylogenetic constraints. Differences in ancestral bipedal taxa within each clade may have resulted in disparate ecomorphological constraints on the evolutionary pathways driving divergence in their quadrupedal descendants.

Ankylosauria is a diverse clade of armoured dinosaurs whose members were important constituents of many Cretaceous faunas. Phylogenetic analyses imply that the clade diverged from its sister taxon, Stegosauria, during the late Early Jurassic, but the fossil records of both clades are sparse until the Late Jurassic (~150 million years ago). Moreover, Ankylosauria is almost entirely restricted to former Laurasian continents, with only a single valid Gondwanan taxon. Spicomellus afer gen. et sp. nov. appears to represent the earliest-known ankylosaur and the first to be named from Africa, from the Middle Jurassic (Bathonian–Callovian) of Morocco, filling an important gap in dinosaur evolution. The specimen consists of a rib with spiked dermal armour fused to its dorsal surface, an unprecedented morphology among extinct and extant vertebrates. The specimen reveals an unrealized morphological diversity of armoured dinosaurs during their early evolution, and implies the presence of an important but undiscovered Gondwanan fossil record. The authors report a fossilized vertebrate rib with spiked dermal armour fused to its dorsal surface from the mid-Jurassic of Morocco, which they interpret as the earliest known ankylosaur.

A new genus and species of non-hadrosaurid hadrosauriform dinosaur, Brighstoneus simmondsi gen. et sp. nov., is described from the Lower Cretaceous Wessex Formation of the Isle of Wight. The new ta...

Neotheropoda represents the main evolutionary radiation of predatory dinosaurs and its oldest records come from Upper Triassic rocks (c. 219 Mya). The Early Jurassic record of Neotheropoda is taxonomically richer and geographically more widespread than that of the Late Triassic. The Lower Jurassic (upper Hettangian–lower Sinemurian) rocks of central England have yielded three neotheropod specimens that have been assigned to two species within the genus Sarcosaurus, S. woodi (type species) and S. andrewsi. These species have received little attention in discussions of the early evolution of Neotheropoda and recently have been considered as nomina dubia. Here, we provide a detailed redescription of one of these specimens (WARMS G667–690) and reassess the taxonomy and phylogenetic relationships of the genus Sarcosaurus. We propose that the three neotheropod specimens from the Early Jurassic of central England represent a single valid species, S. woodi. The second species of the genus, ‘S. andrewsi’, is a subjective junior synonym of the former. A quantitative phylogenetic analysis of early theropods recovered S. woodi as one of the closest sister-taxa to Averostra and provides new information on the sequence of character state transformations in the lead up to the phylogenetic split between Ceratosauria and Tetanurae.

Eurypoda, the major radiation of armoured dinosaurs, comprises the ankylosaurs and their sister group, the stegosaurs. As the earliest-branching major clade of ornithischian dinosaurs, the evolutionary history of Eurypoda is significant for understanding both the palaeobiology of bird-hipped dinosaurs and the composition of middle Mesozoic ecosystems. Eurypodans were diverse and abundant throughout the Late Jurassic and Cretaceous in Laurasia; in contrast, their remains are extremely rare in Gondwana. Herein, we describe a new genus and species of stegosaur from the Middle Jurassic of Morocco, Adratiklit boulahfa. Adratiklit is the first eurypodan from north Africa and the oldest definitive stegosaur from anywhere in the world. The genus is more closely related to the European stegosaurs Dacentrurus and Miragaia than it is to the southern African taxa Kentrosaurus and Paranthodon. Statistically significant correlations between the number of dinosaur-bearing formations, dinosaur-bearing collections, and eurypodan occurrences in Gondwana indicates that their fossil record is biased by both geological and anthropogenic factors. Tantalizing but fragmentary remains and trackways suggest that eurypodan diversity in Gondwana may have been as rich as that of Laurasia, and the prospects for future discoveries of new genera across Gondwana are therefore very good.

Ankylosauria is a diverse clade of armoured dinosaurs whose members were important constituents of many Cretaceous faunas. Phylogenetic analyses imply that the clade diverged from its sister taxon, Stegosauria, during the late Early Jurassic, but the fossil records of both clades are sparse until the Late Jurassic (~150 million years ago). Moreover, Ankylosauria is almost entirely restricted to former Laurasian continents, with only a single valid Gondwanan taxon. Spicomellus afer gen. et sp. nov. appears to represent the earliest-known ankylosaur and the first to be named from Africa, from the Middle Jurassic (Bathonian-Callovian) of Morocco, filling an important gap in dinosaur evolution. The specimen consists of a rib with spiked dermal armour fused to its dorsal surface, an unprecedented morphology among extinct and extant vertebrates. The specimen reveals an unrealized morphological diversity of armoured dinosaurs during their early evolution, and implies the presence of an important but undiscovered Gondwanan fossil record.

The ability of palaeontologists to correctly diagnose and classify new fossil species from incomplete morphological data is fundamental to our understanding of evolution. Different parts of the vertebrate skeleton have different likelihoods of fossil preservation and varying amounts of taxonomic information, which could bias our interpretations of fossil material. Substantial previous research has focused on the diversity and macroevolution of non-avian theropod dinosaurs. Theropods provide a rich dataset for analysis of the interactions between taxonomic diagnosability and fossil preservation. We use specimen data and formal taxonomic diagnoses to create a new metric, the Likelihood of Diagnosis, which quantifies the diagnostic likelihood of fossil species in relation to bone preservation potential. We use this to assess whether a taxonomic identification bias impacts the non-avian theropod fossil record. We find that the patterns of differential species abundance and clade diversity are not a consequence of their relative diagnosability. Although there are other factors that bias the theropod fossil record that are not investigated here, our results suggest that patterns of relative abundance and diversity for theropods might be more representative of Mesozoic ecology than often considered.

The fluvial, overbank, and lacustrine deposits of the Upper Jurassic Morrison Formation of the Western Interior, U.S.A. have been intensively studied due to their diverse and well-preserved dinosaurian fauna, and the presence of economic quantities of uranium and vanadium ores. The formation crops out over 12 degrees of latitude and 1.2 million km2, and is an excellent case study for the examination of paleoecology, community structure, and evolutionary dynamics at a time in Earth's history when the climate was significantly warmer than today. However, paleoecological studies have been hampered by lack of correlation across the formation. Assuming a primarily tectonic control on fluvial architecture, we propose the first chronostratigraphic framework of the formation, which is based on sequence stratigraphy, magnetostratigraphy, and radiometric dating. The formation can be divided into three sequences each represented by a period of degradation followed by aggradation. This chronostratigraphic framework indicates that the formation youngs to the north, and was deposited over about 7 million years during the late Kimmeridgian and Tithonian. This framework provides a foundation for future sedimentological, stratigraphic, and paleobiological studies of the iconic dinosaurian fauna known from the Morrison.

We report a case of a specific osteopathy in the stegosaurian dinosaur Gigantspinosaurus sichuanensis from the Late Jurassic of Zigong City, China. The G. sichuanensis skeleton is very complete. Th...

Erythrosuchidae were large-bodied, quadrupedal, predatory archosauriforms that dominated the hypercarnivorous niche in the aftermath of the Permo-Triassic mass extinction.Garjainia, one of the oldest members of the clade, is known from the late Olenekian of European Russia. The holotype ofGarjainia primacomprises a well-preserved skull, but highly incomplete postcranium. Recent taxonomic reappraisal demonstrates that material from a bone bed found close to the type locality, previously referred to as ‘Vjushkovia triplicostata', is referable toG. prima.At least, seven individuals comprising cranial remains and virtually the entire postcranium are represented, and we describe this material in detail for the first time. An updated phylogenetic analysis confirms previous results that a monophyleticGarjainiais the sister taxon to a clade containingErythrosuchus, ShansisuchusandChalishevia. Muscle scars on many limb elements are clear, allowing reconstruction of the proximal locomotor musculature. We calculate the body mass ofG. primato have been 147–248 kg, similar to that of an adult male lion. Large body size in erythrosuchids may have been attained as part of a trend of increasing body size after the Permo-Triassic mass extinction and allowed erythrosuchids to become the dominant carnivores of the Early and Middle Triassic.

If we imagine walking through Mesozoic lands, we would be able to observe vertebrates with peculiar combinations of morphological traits, some of which would seem to be intermediary to animals seen today. We would witness a terrestrial vertebrate fauna dominated by dinosaurs of various sizes and diversity, accompanied by many other animal groups that often are overlooked. Current research suggests that many of the main vertebrate clades existing today originated or diversified sometime in the Triassic or Early to Middle Jurassic. Herein, we profile some of the major transformations in both terrestrial and aquatic vertebrate evolution during the Mesozoic. We highlight: the appearance of features that allowed sauropod dinosaurs to become the largest animals to ever walk on Earth’s continents, the appearance of herbivory among the usually carnivorous theropod dinosaurs, and we follow the specific changes that led to the evolution of avian flight. Our Mesozoic tour across the globe will allow us to see how different evolutionary forces led to convergent shifts to quadrupedality in ornithischian dinosaurs and to an aquatic lifestyle in turtles, crocodiles, and plesiosaurs. Last, but not least, we examine changes in the Mesozoic fauna linked to the rise of mammals, and the diversification patterns in several clades of fishes after the End-Permian Mass Extinction.

In order for palaeontological data to be informative to ecologists seeking to understand the causes of today's diversity patterns, palaeontologists must demonstrate that actual biodiversity patterns are preserved in our reconstructions of past ecosystems. During the Late Cretaceous, North America was divided into two landmasses, Laramidia and Appalachia. Previous work has suggested strong faunal provinciality on Laramidia at this time, but these arguments are almost entirely qualitative. We quantitatively investigated faunal provinciality in ceratopsid and hadrosaurid dinosaurs using a biogeographic network approach and investigated sampling biases by examining correlations between dinosaur occurrences and collections. We carried out a model-fitting approach using generalized least-squares regression to investigate the sources of sampling bias we identified. We find that while the raw data strongly support faunal provinciality, this result is driven by sampling bias. The data quality of ceratopsids and hadrosaurids is currently too poor to enable fair tests of provincialism, even in this intensively sampled region, which probably represents the best-known Late Cretaceous terrestrial ecosystem on Earth. To accurately reconstruct biodiversity patterns in deep time, future work should focus on smaller scale, higher resolution case studies in which the effects of sampling bias can be better controlled.

Ornithischian dinosaurs were primitively bipedal, but reverted to quadrupedality on at least three (and potentially several more) occasions: in Ceratopsia, Thyreophora and Hadrosauriformes. Each of these reversals was accompanied by anatomical changes to the whole skeleton that enabled the forelimb to function in weight bearing and that also resulted in numerous changes to the hip and hind limb musculature. Each quadrupedal clade acquired a suite of similar biomechanical characters, although they varied in terms of function and in how these character complexes were assembled. Some similar changes occurred in parallel among sauropodomorph dinosaurs as they transitioned from bipedality to quadrupedality. It is unclear why bipedal ornithischians reverted to quadrupedalism, but neither increases in body size nor the acquisition of dermal armour seem to have played a significant role. Increased head size might have influenced the position of the centre of mass and stance in ceratopsians and it is plausible that the evolution of herbivory played an important role in both ornithischians and sauropods, but the latter hypothesis is difficult to test.

The Wealden Supergroup of south-east England has long been of interest to palaeontologists because of its diverse flora and fauna. The Supergroup is Early Cretaceous in age, occupying the time period immediately after the enigmatic end-Jurassic extinction. Wealden faunas therefore have the potential to be informative about the tempo and mode of post-extinction recovery, but due to lack of exposure in this densely populated part of southern England, are difficult to sample. In the summer of 2012, a number of ex situ fossiliferous blocks of sandstone, siltstone and limestone were discovered from building excavations at Ardingly College, near Haywards Heath in West Sussex. The sedimentology of the blocks indicates that they are from the Valanginian Hastings Group, and that Ardingly College is underlain by the Grinstead Clay Formation, rather than the Ardingly Sandstone Member. The blocks contain a diverse invertebrate fauna and flora, as well as vertebrate remains, which are found in a distinct sandstone horizon that probably represents the Top Lower Tunbridge Wells pebble bed. A tooth from an ornithschian dinosaur cannot be referred to any of the ornithischian taxa known from the Wealden Supergroup, and therefore represents a new taxon. Teeth of the crocodilian Theriosuchus extend the known range of this taxon in the Wealden, while teeth of an ornithocheird pterosaur confirm the presence of these animals in the skies above the Wealden sub-basins. Fusainized plant remains and the wing-case of a cupedid beatle indicate that wildfire was a ubiquitous feature of the Weald Sub-basin during the Valanginian.

Erythrosuchidae are a globally distributed and important group of apex predators that occupied Early and Middle Triassic terrestrial ecosystems following the Permo-Triassic mass extinction. The stratigraphically oldest known genus of Erythrosuchidae is Garjainia Ochev, 1958, which is known from the late Early Triassic (late Olenekian) of European Russia and South Africa. Two species of Garjainia have been reported from Russia: the type species, Garjainia prima Ochev, 1958, and ‘Vjushkovia triplicostata’ von Huene, 1960, which has been referred to Garjainia as either congeneric (Garjainia triplicostata) or conspecific (G. prima). The holotype of G. prima has received relatively extensive study, but little work has been conducted on type or referred material attributed to ‘V. triplicostata’. However, this material includes well-preserved fossils representing all parts of the skeleton and comprises seven individuals. Here, we provide a comprehensive description and review of the cranial anatomy of material attributed to ‘V. triplicostata’, and draw comparisons with G. prima. We conclude that the two Russian taxa are indeed conspecific, and that minor differences between them result from a combination of preservation or intraspecific variation. Our reassessment therefore provides additional information on the cranial anatomy of G. prima. Moreover, we quantify relative head size in erythrosuchids and other early archosauromorphs in an explicit phylogenetic context for the first time. Our results show that erythrosuchids do indeed appear to have disproportionately large skulls, but that this is also true for other early archosauriforms (i.e. proterosuchids), and may reflect the invasion of hypercarnivorous niches by these groups following the Permo-Triassic extinction. Fil: Butler, Richard J.. University of Birmingham; Reino Unido Fil: Sennikov, Andrey G.. Paleontological Institute of Russian Academy of Sciences; Rusia. Kazan Federal University; Rusia Fil: Dunne, Emma M.. University of Birmingham; Reino Unido Fil: Ezcurra, Martin Daniel. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina. University of Birmingham; Reino Unido Fil: Hedrick, Brandon P.. University of Oxford; Reino Unido Fil: Maidment, Susannah C. R.. Natural History Museum; Reino Unido. University of Birmingham; Reino Unido Fil: Meade, Luke E.. University of Birmingham; Reino Unido Fil: Raven, Thomas J.. University of Brighton; Reino Unido. Natural History Museum; Reino Unido Fil: Gower, David J.. Natural History Museum; Reino Unido

Ankylosaurs, dinosaurs possessing extensive body armor, were significant components of terrestrial ecosystems from the Middle Jurassic–latest Cretaceous. They diversified during the Early Cretaceou...

The first African dinosaur to be discovered, Paranthodon africanus was found in 1845 in the Lower Cretaceous of South Africa. Taxonomically assigned to numerous groups since discovery, in 1981 it was described as a stegosaur, a group of armoured ornithischian dinosaurs characterised by bizarre plates and spines extending from the neck to the tail. This assignment has been subsequently accepted. The type material consists of a premaxilla, maxilla, a nasal, and a vertebra, and contains no synapomorphies of Stegosauria. Several features of the maxilla and dentition are reminiscent of Ankylosauria, the sister-taxon to Stegosauria, and the premaxilla appears superficially similar to that of some ornithopods. The vertebral material has never been described, and since the last description of the specimen, there have been numerous discoveries of thyreophoran material potentially pertinent to establishing the taxonomic assignment of the specimen. An investigation of the taxonomic and systematic position of Paranthodon is therefore warranted. This study provides a detailed re-description, including the first description of the vertebra. Numerous phylogenetic analyses demonstrate that the systematic position of Paranthodon is highly labile and subject to change depending on which exemplifier for the clade Stegosauria is used. The results indicate that the use of a basal exemplifier may not result in the correct phylogenetic position of a taxon being recovered if the taxon displays character states more derived than those of the basal exemplifier, and we recommend the use, minimally, of one basal and one derived exemplifier per clade. Paranthodon is most robustly recovered as a stegosaur in our analyses, meaning it is one of the youngest and southernmost stegosaurs.

The stegosaurs are some of the most easily recognizable dinosaurs, but are surprisingly rare as fossils. Consequently much remains unknown about their palaeobiology, and every new stegosaurian find contributes to our understanding of the evolution of the clade. Since the last attempt to examine the evolutionary relationships of Stegosauria, new specimens have come to light, including the most complete individual of Stegosaurus ever found, new taxa have been described and, perhaps most importantly, new methods for analysis of cladistic datasets have been produced. In the light of these new data and technological advances, the phylogenetic relationships of the stegosaurs and basal armoured dinosaurs are investigated. The inclusion of continuous data results in much better resolution than was previously obtained, and the resulting single most parsimonious tree supports re‐erection of the genera Miragaia and Hesperosaurus, which had previously been synonymized with Dacentrurus and Stegosaurus respectively. The recently described genus Alcovasaurus is resolved as a basal thyreophoran, but this is most likely a consequence of a very high degree of missing data and the questionable ontogenetic stage of the specimen. Examination of the effects of continuous data on the analysis suggest that while it contains a phylogenetic signal congruent with that of discrete data and provides better resolution than discrete data alone, it can affect topologies in unpredictable ways, particularly in areas of the tree where there are large amounts of missing data. The phylogeny presented here will form the basis for future work on the palaeobiology of the plated dinosaurs.

The first African dinosaur to be discovered,Paranthodon africanuswas found in 1845 in the Lower Cretaceous of South Africa. Taxonomically assigned to numerous groups since discovery, in 1981 it was described as a stegosaur, a group of armoured ornithischian dinosaurs characterised by bizarre plates and spines extending from the neck to the tail. This assignment has been subsequently accepted. The type material consists of a premaxilla, maxilla, a nasal, and a vertebra, and contains no synapomorphies of Stegosauria. Several features of the maxilla and dentition are reminiscent of Ankylosauria, the sister-taxon to Stegosauria, and the premaxilla appears superficially similar to that of some ornithopods. The vertebral material has never been described, and since the last description of the specimen, there have been numerous discoveries of thyreophoran material potentially pertinent to establishing the taxonomic assignment of the specimen. An investigation of the taxonomic and systematic position ofParanthodonis therefore warranted. This study provides a detailed re-description, including the first description of the vertebra. Numerous phylogenetic analyses demonstrate that the systematic position ofParanthodonis highly labile and subject to change depending on which exemplifier for the clade Stegosauria is used. The results indicate that the use of a basal exemplifier may not result in the correct phylogenetic position of a taxon being recovered if the taxon displays character states more derived than those of the basal exemplifier, and we recommend the use, minimally, of one basal and one derived exemplifier per clade.Paranthodonis most robustly recovered as a stegosaur in our analyses, meaning it is one of the youngest and southernmost stegosaurs.

The moment arm of a muscle defines its leverage around a given joint. In a clinical setting, the quantification of muscle moment arms is an important means of establishing the ‘healthy’ functioning of a muscle and in identifying and treating musculoskeletal abnormalities. Elsewhere in modern animal taxa, moment arm studies aim to illuminate adaptions of the musculoskeletal system towards particular locomotor or feeding behaviors. In the absence of kinematic data, paleontologists have likewise relied upon estimated muscle moment arms as a means of reconstructing musculoskeletal function and biomechanical performance in fossil species. With the application of ‘virtual paleontological’ techniques, it is possible to generate increasingly detailed musculoskeletal models of extinct taxa. However, the steps taken to derive such models of complex systems are seldom reported in detail. Here we present a case study for calculating three-dimensional muscle moment arms using Stegosaurus stenops Marsh, 1887 to highlight both the potential and the limitations of this approach in vertebrate paleontology. We find the technique to be mostly insensitive to choices in muscle modeling parameters (particularly relative to other sources of uncertainty in paleontological studies), although exceptions do exist. Of more concern is the current lack of consensus on what functional signals, if any, are contained within moment arm data derived from extant species. Until a correlation between muscle moment arm and function can be broadly identified across a range of modern taxa, the interpretation of moment arms calculated for extinct taxa should be approached with caution.

The stegosaurs are some of the most easily recognisable dinosaurs, but they are surprisingly rare as fossils. Consequently much remains unknown about their palaeobiology, and every new stegosaurian find contr ibutes to understanding the evolution of the clade. Since the last attempt to examine the evolutionary relationships of Stegosauria, new specimens have come to light, including the most complete individual of Stegosaurus ever found, new taxa have been desc ribed and, perhaps most importantly, new methods for analysis of cladistic datasets have been produced. In the light of these new data and technological advances, the phylogenetic relationships of the stegosaurs and basal armoured dinosaurs are investigate d. The inclusion of continuous data results in much better resolution than was previously obtained, and the resulting single most parsimonious tree supports re - erection of the genera Miragaia and Hesperosaurus , which had previously been synonymized with Dacentrurus and Stegosaurus respectively. The recently described genus Alcovasaurus is resolved as a basal thyreophoran, but this is likely to be due to a combination of a very high degree of missing data and the questionable ontogenetic stage of the speci men. Examination of the effects of continuous data on the analysis suggest that while it contains a phylogenetic signal congruent with that of discrete data and provides better resolution than discrete data alone, it can affect t opologies in unpredictable ways, particularly in areas of the tree where there are large amounts of missing data. The phylogeny presented here will form the basis for future work on the palaeobiology of the plated dinosaurs.

The terrestrial deposits of the Upper Jurassic Morrison Formation, western USA, have been intensively studied since the discovery of their well-preserved and diverse dinosaurian fauna. The broad geographic area and large time interval represented by the formation makes it an ideal candidate for the study of biodiversity patterns, population ecology, and morphological evolution through time, with environment and with latitude, but attempts to use it for such studies have been hampered by a lack of long-range correlation. We collected magnetostratigraphic samples near Dinosaur National Monument, Utah, in order to examine the magnetostratigraphy of this heavily studied area, and assess the prospects of using magnetostratigraphy as a correlative tool in the Morrison Formation. A magnetic reversal immediately above the quarry sand at Dinosaur National Monument, in combination with an existing radiometric date, allows us to constrain the age of the historic dinosaur quarry to 150.91–150.04 million years. Along with radiometric dating, magnetostratigraphy is useful for tightly constraining the ages of dinosaur quarries to allow faunal comparisons through space and time. Problems encountered during sampling might be alleviated by collecting hand specimens in Morrison mudstones.

The Upper Jurassic Morrison Formation of the western United States preserves one of the best-known Mesozoic paleoecosystems worldwide. The formation crops out over an area from New Mexico and Oklahoma to Montana and Utah and encompasses a time span of approximately eight million years. Recent studies indicate a high diversity of gigantic, herbivorous sauropod dinosaurs, but the geographic and temporal distributions of species or even genera of these animals remain poorly understood. In particular, sauropod specimens from northern outcrops of the formation have rarely been studied in detail, and temporal relationships among sites are imprecise. Here, we reassess the taxonomic diversity of the sauropods from a historic Carnegie Museum locality in northern Wyoming. Previous referrals of material to the well-known diplodocid genera Apatosaurus and Diplodocus cannot be confidently confirmed; instead, all these specimens more likely represent elements from the recently recognized Galeamopus. Specimens previously assigned to Camarasaurus and Haplocanthosaurus could not be referred to these genera based on apomorphies, due to a lack of detailed knowledge concerning the genus- and species-level taxonomy of these sauropods. Our findings imply that many referrals of incomplete diplodocid skeletons to Apatosaurus and Diplodocus must be reassessed. These reassessments are particularly important with regard to specimens from northern localities of the Morrison Formation, as it is becoming increasingly evident that diplodocids from this area were distinct from better-known, more southerly taxa. This geographic segregation does not seem to apply to nondiplodocid sauropods; however, these taxa are also in need of systematic revision, which may reveal species-level patterns similar to those observed in Diplodocidae.

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionary history. The osteological and myological changes associated with these transitions have only recently been documented, and the biomechanical consequences of these changes remain to be examined. Here, we review previous approaches to understanding locomotion in extinct animals, which can be broadly split into form–function approaches using analogy based on extant animals, limb-bone scaling, and computational approaches. We then carry out the first systematic attempt to quantify changes in locomotor muscle function in bipedal and quadrupedal ornithischian dinosaurs. Using three-dimensional computational modelling of the major pelvic locomotor muscle moment arms, we examine similarities and differences among individual taxa, between quadrupedal and bipedal taxa, and among taxa representing the three major ornithischian lineages (Thyreophora, Ornithopoda, Marginocephalia). Our results suggest that the ceratopsid Chasmosaurus and the ornithopod Hypsilophodon have relatively low moment arms for most muscles and most functions, perhaps suggesting poor locomotor performance in these taxa. Quadrupeds have higher abductor moment arms than bipeds, which we suggest is due to the overall wider bodies of the quadrupeds modelled. A peak in extensor moment arms at more extended hip angles and lower medial rotator moment arms in quadrupeds than in bipeds may be due to a more columnar hindlimb and loss of medial rotation as a form of lateral limb support in quadrupeds. We are not able to identify trends in moment arm evolution across Ornithischia as a whole, suggesting that the bipedal ancestry of ornithischians did not constrain the development of quadrupedal locomotion via a limited number of functional pathways. Functional anatomy appears to have had a greater effect on moment arms than phylogeny, and the differences identified between individual taxa and individual clades may relate to differences in locomotor performance required for living in different environments or for clade-specific behaviours.

Convergent morphologies are thought to indicate functional similarity, arising because of a limited number of evolutionary or developmental pathways. Extant taxa displaying convergent morphologies are used as analogues to assess function in extinct taxa with similar characteristics. However, functional studies of extant taxa have shown that functional similarity can arise from differing morphologies, calling into question the paradigm that form and function are closely related. We test the hypothesis that convergent skeletal morphology indicates functional similarity in the fossil record using ornithischian dinosaurs. The rare transition from bipedality to quadrupedality occurred at least three times independently in this clade, resulting in a suite of convergent osteological characteristics. We use homology rather than analogy to provide an independent line of evidence about function, reconstructing soft tissues using the extant phylogenetic bracket and applying biomechanical concepts to produce qualitative assessments of muscle leverage. We also optimize character changes to investigate the sequence of character acquisition. Different lineages of quadrupedal ornithischian dinosaur stood and walked differently from each other, falsifying the hypothesis that osteological convergence indicates functional similarity. The acquisition of features correlated with quadrupedalism generally occurs in the same order in each clade, suggesting underlying developmental mechanisms that act as evolutionary constraints.

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.

The earliest ornithischian dinosaurs were small bipeds that possessed the derived, retroverted pubis that is synapomorphic for the clade. Their forelimbs were modified for grasping. Subsequently, ornithischians radiated into numerous body shapes and sizes. Three lineages independently evolved large size and quadrupedality, requiring profound changes to the osteology and myology of the locomotor apparatus. Using comparisons with extinct archosaurian outgroups, as well as extant birds and crocodilians, we reconstruct basal ornithischian limb musculature in order to determine the sequence of musculoskeletal changes that occurred prior to and during the early evolution of Ornithischia. The musculoskeletal anatomy of basal ornithischian forelimbs was probably more similar to that of the crown group archosaur common ancestor and crocodilians than it was to birds. Reduction or loss of the clavicle resulted in migration of the clavicular deltoid onto the proximal scapula and development of a distinct acr...

Sexual dimorphism, the condition whereby males and females differ from one another physically, is one of the most fundamental aspects of the biology of any animal. However, sexually dimorphic characters can be subtle and are mainly related to soft tissue anatomy. They are, therefore, difficult to identify reliably in the fossil record particularly when dealing with small sample sizes and osteology alone. The first geometric morphometric analysis of dimorphism in a thyreophoran (armored) dinosaur shows that the femora of the stegosaurian dinosaur Kentrosaurus aethiopicus (Upper Jurassic, Tanzania) bear a statistically significant shape difference of the proximal end, which is independent of size and is therefore proposed to be a sexual difference. Although the disarticulated nature of the material means that intraspecific variation in other skeletal elements, such as the enigmatic dermal armor, cannot be identified as sexual dimorphism at this time, this study provides a methodology for further wo...

The first partial skeleton of a stegosaurian dinosaur was discovered in a brick pit in Swindon, UK in 1874. Since then, numerous stegosaurian remains have been discovered from Europe, North America, Africa and Asia, and continue to be discovered regularly. Stegosaurs are known from the Middle Jurassic to the Early Cretaceous; no definitive evidence of the clade is known from younger deposits. New discoveries are improving our understanding of stegosaur biology and showing that stegosaurs were more morphologically diverse than was previously realized. A new phylogeny, which includes all valid stegosaurian taxa, largely agrees with previous studies and shows the European Dacentrurinae was sister taxon to Stegosaurus. Poor resolution at the base of Stegosauria is probably due to the fragmentary nature of many of the Chinese taxa.

The palpebral is a small ossification that projects across the orbit in some ornithischian dinosaurs and its presence is considered a synapomorphy of the clade. By contrast, other ornithischians lack the palpebral but possess accessory ossifications, commonly termed supraorbitals, which form the dorsal margin of the orbit. The homology of the ornithischian palpebral to one or more of the supraorbitals is widely accepted in the literature, but this homology has never been explicitly tested and no hypotheses have been proposed regarding the function of the palpebral or why it was incorporated into the orbital margin. As homology is synonymous with synapomorphy, incorrect homology statements can lead to incorrect hypotheses of relationships being obtained during cladistic analysis. The primary and secondary homologies of the ornithischian palpebral and the anterior supraorbital of more derived members of the major ornithischian clades are tested and we demonstrate that these homology hypotheses can be accepted. Osteological correlates indicate that the palpebral supported a layer of connective tissue that roofed the orbit; ossification of this connective tissue resulted in the incorporation of the palpebral into the skull roof and gave rise to additional supraorbital elements, which are neomorphic ossifications. Large-scale structural changes in the ornithischian skull, including dermal ossifications associated with display or defence and the development of complex feeding mechanisms, may have led to the incorporation of the palpebral into the skull roof.

Stegosauria is a clade of ornithischian dinosaurs characterized by a bizarre array of dermal armor that extends from the neck to the end of the tail. Two genera of stegosaur are currently recognised from North America: the well-known Stegosaurus stenops and the much rarer Hesperosaurus mjosi. A new specimen of Hesperosaurus mjosi was discovered in some of the most northerly outcrops of the Upper Jurassic Morrison Formation near Livingston, Montana. The new specimen includes cranial, vertebral, and appendicular material as well as a dermal plate, and the excellent state of preservation of the palate reveals new anatomical information about this region in stegosaurs. Histological examination of the tibia indicates that the individual was not skeletally mature at time of death. Comparison with previously studied Stegosaurus and Hesperosaurus individuals indicates that Hesperosaurus mjosi may have been a smaller species than Stegosaurus stenops. Physiological processes scale with body mass, M, accordi...

Synopsis Stegosauria is a clade of ornithischian dinosaurs characterised by a bizarre array of dermal armour extending, in two parasagittal rows, from the cervical region to the end of the tail. Although Stegosaurus is one of the most familiar of all dinosaurs, little is known regarding the evolutionary history of this clade. Alpha‐level taxonomic revision of all proposed stegosaur taxa shows that 11 species of stego‐saur can be regarded as valid on the basis of autapomorphies. These are: Dacentrurus armatus and Loricatosaurus priscus (gen. nov.) from Europe; Kentrosaurus aethiopicus and Paranthodon afric‐anus from Africa; Tuojiangosaurus multispinus, Chungkingosaurus jiangbeiensis, Huayangosaurus taibaii, Gigantspinosaurus sichuanensis and Stegosaurus homheni (comb. nov.) from China; and Stegosaurus mjosi (comb. nov.) and Stegosaurus armatus from North America. A cladistic analysis of Stegosauria (the first to be based upon direct observation of all relevant specimens) is presented, which indicates that ...

The early Mesozoic fossil fauna collected from the Lower Lufeng Formation of Yunnan Province, China, has attracted considerable interest and attention since its discovery in the late 1930s. Its importance reflected a combination of its comparatively remote geographical position and, more particularly, the similarities of its fauna compared with approximately contemporary discoveries from Europe, North and South America, and southern Africa. The fragmentary and poorly preserved Lufeng ornithischian dinosaur Tatisaurus oehleri was described in 1965 and proved taxonomically and systematically enigmatic from the start. Originally assigned, with some noted ambivalence, to the basal ('primitive') group of ornithischians known as hypsilophodontids, since 1965 Tatisaurus has been variously ignored, assigned to a more rigorously defined Hypsilophodontidae, referred to both of the armoured (thyreophoran) ornithischian dinosaur clades (Stegosauria and Ankylosauria), or referred to a more basal position within the thyreophoran lineage. In 1996 the holotype of Tatisaurus was renamed Scelidosaurus oehleri, and the genus Scelidosaurus was proposed as an index fossil of the 'Scelidosaurus biochron' with the potential to be used for the global stratigraphic correlation of Early Jurassic (early Sinemurian) rocks. Because of this chequered history Tatisaurus oehleriSimmons, 1965 has been re-examined and is redescribed so that its taxonomic status and systematic position could be reassessed. Tatisaurus is identified as a basal thyreophoran (armoured ornithischian dinosaur); there is no basis for amalgamating it in synonymy with the genus Scelidosaurus, and the proposed creation of a 'Scelidosaurus biochron' for the purposes of biostratigraphic correlation of Lower Jurassic outcrops has no utility whatever. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 150, 865–874.

In a number of recent studies we used computer modeling to investigate the evolution of muscle leverage (moment arms) and function in extant and extinct archosaur lineages (crocodilians, dinosaurs including birds and pterosaurs). These studies sought to quantify the level of disparity and convergence in muscle moment arms during the evolution of bipedal and quadrupedal posture in various independent archosaur lineages, and in doing so further our understanding of changes in anatomy, locomotion and ecology during the group’s >250 million year evolutionary history. Subsequent work by others has led us to re-evaluate our models, which revealed a methodological error that impacted on the results obtained from the abduction–adduction and long-axis rotation moment arms in our published studies. In this paper we present corrected abduction–adduction and long axis rotation moment arms for all our models, and evaluate the impact of this new data on the conclusions of our previous studies. We find that, in general, our newly corrected data differed only slightly from that previously published, with very few qualitative changes in muscle moments (e.g., muscles originally identified as abductors remained abductors). As a result the majority of our previous conclusions regarding the functional evolution of key muscles in these archosaur groups are upheld.

Although Stegosaurus is one of the most iconic dinosaurs, well-preserved fossils are rare and as a consequence there is still much that remains unknown about the taxon. A new, exceptionally complete individual affords the opportunity to describe the anatomy of Stegosaurus in detail for the first time in over a century, and enables additional comparisons with other stegosaurian dinosaurs. The new specimen is from the Red Canyon Ranch Quarry, near Shell Wyoming, and appears to have been so well preserved because it was buried rapidly in a pond or body of standing water immediately after death. The quarry is probably located in the middle part of the Morrison Formation, which is believed to be Tithonian in age in this area. The specimen is referable to Stegosaurus stenops based on the possession of an edentulous anterior portion of the dentary and elevated postzygapophyses on the cervical vertebrae. New information provided by the specimen concerns the morphology of the vertebrae, the iliosacral block and dermal armor. Several aspects of its morphology indicate the individual was not fully skeletally mature at the time of death, corroborating a previous histological study.

Exceptionally preserved organic remains are known throughout the vertebrate fossil record, and recently, evidence has emerged that such soft tissue might contain original components. We examined samples from eight Cretaceous dinosaur bones using nano-analytical techniques; the bones are not exceptionally preserved and show no external indication of soft tissue. In one sample, we observe structures consistent with endogenous collagen fibre remains displaying ∼67 nm banding, indicating the possible preservation of the original quaternary structure. Using ToF-SIMS, we identify amino-acid fragments typical of collagen fibrils. Furthermore, we observe structures consistent with putative erythrocyte remains that exhibit mass spectra similar to emu whole blood. Using advanced material characterization approaches, we find that these putative biological structures can be well preserved over geological timescales, and their preservation is more common than previously thought. The preservation of protein over geological timescales offers the opportunity to investigate relationships, physiology and behaviour of long extinct animals., Soft tissue from vertebrate fossils has previously been documented, but only in exceptionally preserved specimens. Here, Bertazzo et al. describe structures consistent with collagen fibres and red blood cells from eight Cretaceous dinosaur bones, none of which are exceptionally preserved.