Basal Tetanuran Research Articles

Modified skulls but conservative brains? The palaeoneurology and endocranial anatomy of baryonychine dinosaurs (Theropoda: Spinosauridae).

The digital reconstruction of neurocranial endocasts has elucidated the gross brain structure and potential ecological attributes of many fossil taxa, including Irritator, a spinosaurine spinosaurid from the "mid" Cretaceous (Aptian) of Brazil. With unexceptional hearing capabilities, this taxon was inferred to integrate rapid and controlled pitch-down movements of the head that perhaps aided in the predation of small and agile prey such as fish. However, the neuroanatomy of baryonychine spinosaurids remains to be described, and potentially informs on the condition of early spinosaurids. Using micro-computed tomographic scanning (μCT), we reconstruct the braincase endocasts of Baryonyx walkeri and Ceratosuchops inferodios from the Wealden Supergroup (Lower Cretaceous) of England. We show that the gross endocranial morphology is similar to other non-maniraptoriform theropods, and corroborates previous observations of overall endocranial conservatism amongst more basal theropods. Several differences of unknown taxonomic utility are noted between the pair. Baryonychine neurosensory capabilities include low-frequency hearing and unexceptional olfaction, whilst the differing morphology of the floccular lobe tentatively suggests less developed gaze stabilisation mechanisms relative to spinosaurines. Given the morphological similarities observed with other basal tetanurans, baryonychines likely possessed comparable behavioural sophistication, suggesting that the transition from terrestrial hypercarnivorous ancestors to semi-aquatic "generalists" during the evolution of Spinosauridae did not require substantial modification of the brain and sensory systems.

Theropod and ornithischian dinosaur track assemblages from Middle to ?Late Jurassic deposits of the Central High Atlas, Morocco

ABSTRACT Tridactyl theropod and ornithischian dinosaur tracks and trackways from Imilchil and Isli formations (Middle–?Late Jurassic, Bajocian–?) of the central High Atlas region (Morocco) are described. The Imilchil Formationconsists of brackish marine-continental deposits, and the Isli Formation is a continental red-bed succession. Considering numerous new footprint discoveries, including recently described Polyonyx sauropod trackways, tridactyl dinosaur tracks from the Imilchil-Tounfite region are revised. Dominating are theropod footprints resembling the ichnogenus Changpeipus and known from Lower-Middle Jurassic deposits of China. Other theropod ichnotaxa are Trisauropodiscus isp., cf. Wildeichnus isp.,Carmelopodus isp., Megalosauripus isp., Kayentapus isp. and an indeterminate small grallatorid. Ornithischians are represented by small indeterminate ornithischian tracks, large ornithischian footprints cf. Stegopodus isp. and a large indeterminate ornithopod form. Makers of the small theropod trackswere small coelurosaurs or basal tetanurans, larger forms belong toceratosaurs, megalosauroids, allosauroids or tyrannosauroids. Ornithischian tracks suggest dryomorphs, iguanodontians and thyreophorans as producers. Together with crocodylomorph and pterosaur tracks, invertebrate traces and plants, the Imilchil and Isli formations document a flourishing ecosystem and dinosaur habitat. Remarkable is the presence of Changpeipus theropod tracks known from abundant occurrences in East Asia. This suggests an exchange of dinosaur faunas between this region and northern Africa during the Middle Jurassic.

A new theropod dinosaur from the early cretaceous (Barremian) of Cabo Espichel, Portugal: Implications for spinosaurid evolution.

Spinosaurids are some of the most enigmatic Mesozoic theropod dinosaurs due to their unique adaptations to aquatic environments and their relative scarcity. Their taxonomy has proven to be especially problematic. Recent discoveries from Western Europe in general, specifically Iberia, provide some of the best specimens for the understanding of their phylogeny, leading to the description of the spinosaurid Vallibonavenatrix cani and the recognition of the Iberian dinosaur Camarillasaurus cirugedae as one of them. Portuguese associated spinosaurid remains (ML1190) from the Papo Seco Formation (early Barremian) were previously assigned to Baryonyx walkeri but new material recovered in 2020 along with new phylogenetic analyses suggests a different phylogenetic placement, making their revision necessary. Here we show that these remains are not attributable to Baryonyx walkeri, but to a new genus and species, Iberospinus natarioi, gen. et sp. nov. The new taxon is characterized by the presence of a single Meckelian foramen in the Meckelian sulcus, a straight profile of the ventral surface of the dentary and a distal thickening of the acromion process of the pubis between other characters. Iberospinus natarioi is recovered as a sister taxon of the clade formed by Baryonyx and Suchomimus, and outside Spinosaurinae when Vallibonaventrix cani is excluded from the analysis. The description of this taxon reinforces Iberia as a hotspot for spinosaur biodiversity, with several endemic taxa for the region. As expected for the clade, the dentary displays a highly vascularized neurovascular network. The morphometric analysis of parts of the skeleton (pedal phalanx and caudal vertebrae, among others) shows an intermediate condition between basal tetanurans and spinosaurines.

Fast-running theropods tracks from the Early Cretaceous of La Rioja, Spain

Theropod behaviour and biodynamics are intriguing questions that paleontology has been trying to resolve for a long time. The lack of extant groups with similar bipedalism has made it hard to answer some of the questions on the matter, yet theoretical biomechanical models have shed some light on the question of how fast theropods could run and what kind of movement they showed. The study of dinosaur tracks can help answer some of these questions due to the very nature of tracks as a product of the interaction of these animals with the environment. Two trackways belonging to fast-running theropods from the Lower Cretaceous Enciso Group of Igea (La Rioja) are presented here and compared with other fast-running theropod trackways published to date. The Lower Cretaceous Iberian fossil record and some features present in these footprints and trackways suggest a basal tetanuran, probably a carcharodontosaurid or spinosaurid, as a plausible trackmaker. Speed analysis shows that these trackways, with speed ranges of 6.5–10.3 and 8.8–12.4 ms−1, testify to some of the top speeds ever calculated for theropod tracks, shedding light on the question of dinosaur biodynamics and how these animals moved.

A brackish to non-marine aquatic and terrestrial fossil assemblage with vertebrates from the lower Coniacian (Upper Cretaceous) Gosau Group of the Tiefengraben locality near St. Wolfgang im Salzkammergut, Austria

The Turonian–Coniacian continental fossil record in Europe is scarce. Here we present a new fossil assemblage of early Coniacian age that was systematically collected from the coal-bearing Gosau Group of the Tiefengraben locality near St. Wolfgang, Austria. The diverse assemblage is composed of at least 60 taxa including sporomorphs and Normapolles-related pollen, seeds and leaves of angiosperms and gymnosperms, charophytes, gastropods, bivalves, ostracods, termites, fishes, crocodiles and dinosaurs. Concerning charophytes, ostracods, gastropods, crocodiles and dinosaurs, the discovered specimens either extend the temporal and spatial range of specific groups (in some cases as possible relict forms) or suggest the occurrence of new taxa. The discovered remains of algae, molluscs, ostracods, calcareous nannofossils and lepisosteid fish represent a mixed faunal assemblage from different palaeohabitats, from marginal marine to low salinity and freshwater or terrestrial environments. As Normapolles-related angiosperm plants dominate the flora with a relatively high number of dentate leaves, a slightly cooler microenvironment compared to other Turonian–Coniacian Central European localities is indicated. The characteristically grooved crocodylian teeth of Tiefengraben differ from the previously known Upper Cretaceous European crocodyliform teeth and suggest a more diverse crocodyliform fauna in the region. Dinosaurs are represented by teeth of at least three different theropods, the largest of which is referred here to as basal tetanurans. The fossil assemblage of this early Gosau Group occurrence is of great importance for our understanding of the continental floristic and faunistic composition of the western Tethyan archipelago during the Cenomanian–Campanian gap.

Notes on the cheek region of the Late Jurassic theropod dinosaur Allosaurus.

Allosaurus, from the Late Jurassic of North America and Europe, is a model taxon for Jurassic basal tetanuran theropod dinosaurs. It has achieved an almost iconic status due to its early discovery in the late, 19th century, and due to the abundance of material from the Morrison Formation of the western U.S.A., making Allosaurus one of the best-known theropod taxa. Despite this, various aspects of the cranial anatomy of Allosaurus are surprisingly poorly understood. Here, we discuss the osteology of the cheek region, comprised by the jugal, maxilla, and lacrimal. This region of the skull is of importance for Allosaurus taxonomy and phylogeny, particularly because Allosaurus has traditionally been reconstructed with an unusual cheek configuration, and because the European species Allosaurus europaeus has been said to be different from North American material in the configuration of these bones. Based on re-examination of articulated and disarticulated material from a number of repositories, we show that the jugal participates in the antorbital fenestra, contradicting the common interpretation. The jugal laterally overlies the lacrimal, and forms an extended antorbital fossa with this bone. Furthermore, we document previously unrecorded pneumatic features of the jugal of Allosaurus.

Small theropod teeth (Dinosauria) from the Upper Jurassic Qigu Formation of the southern Junggar Basin, NW China

Two small theropod teeth from the Qigu Formation of Liuhuanggou, Xinjiang Uygur Autonomous Region, People's Republic of China are described. They are compared to hitherto known finds of theropod teeth from both the northern and southern Junggar Basin. It is concluded that they cannot be confidently referred to any morphotype hitherto known from the Upper Jurassic of the Junggar Basin. Although the teeth agree well in principal measurements with material previously described and referred to basal tetanurans or basal tyrannosauroids (morphotype 4 of Han et al. 2011), they are both considerably smaller and show the distinctive feature of a lingually displaced mesial carina. The latter feature is shared with Dromaeosaurus albertensis and an unnamed dromaeosaur from the Lower Cretaceous of Spain and Denmark. It is therefore concluded that the material possibly represents a dromaeosaurid close to these taxa. It is the first record of putative dromaeosaurid theropods from the southern Junggar Basin. As dromaeosaurids have been previously announced from the Upper Jurassic of the northern Junggar Basin, this further underlines general similarity in the Upper Jurassic dinosaurian faunas between these areas.

The oldest ceratosaurian (Dinosauria: Theropoda), from the Lower Jurassic of Italy, sheds light on the evolution of the three-fingered hand of birds.

The homology of the tridactyl hand of birds is a still debated subject, with both paleontological and developmental evidence used in support of alternative identity patterns in the avian fingers. With its simplified phalangeal morphology, the Late Jurassic ceratosaurian Limusaurus has been argued to support a II–III–IV digital identity in birds and a complex pattern of homeotic transformations in three-fingered (tetanuran) theropods. We report a new large-bodied theropod, Saltriovenator zanellai gen. et sp. nov., based on a partial skeleton from the marine Saltrio Formation (Sinemurian, lowermost Jurassic) of Lombardy (Northern Italy). Taphonomical analyses show bone bioerosion by marine invertebrates (first record for dinosaurian remains) and suggest a complex history for the carcass before being deposited on a well-oxygenated and well-illuminated sea bottom. Saltriovenator shows a mosaic of features seen in four-fingered theropods and in basal tetanurans. Phylogenetic analysis supports sister taxon relationships between the new Italian theropod and the younger Early Jurassic Berberosaurus from Morocco, in a lineage which is the basalmost of Ceratosauria. Compared to the atrophied hand of later members of Ceratosauria, Saltriovenator demonstrates that a fully functional hand, well-adapted for struggling and grasping, was primitively present in ceratosaurians. Ancestral state reconstruction along the avian stem supports 2-3-4-1-X and 2-3-4-0-X as the manual phalangeal formulae at the roots of Ceratosauria and Tetanurae, confirming the I–II–III pattern in the homology of the avian fingers. Accordingly, the peculiar hand of Limusaurus represents a derived condition restricted to late-diverging ceratosaurians and cannot help in elucidating the origin of the three-fingered condition of tetanurans. The evolution of the tridactyl hand of birds is explained by step-wise lateral simplification among non-tetanuran theropod dinosaurs, followed by a single primary axis shift from digit position 4 to 3 at the root of Tetanurae once the fourth finger was completely lost, which allowed independent losses of the vestigial fourth metacarpal among allosaurians, tyrannosauroids, and maniraptoromorphs. With an estimated body length of 7.5 m, Saltriovenator is the largest and most robust theropod from the Early Jurassic, pre-dating the occurrence in theropods of a body mass approaching 1,000 Kg by over 25 My. The radiation of larger and relatively stockier averostran theropods earlier than previously known may represent one of the factors that ignited the trend toward gigantism in Early Jurassic sauropods.

The largest European theropod dinosaurs: remains of a gigantic megalosaurid and giant theropod tracks from the Kimmeridgian of Asturias, Spain.

The Kimmeridgian Vega, Tereñes and Lastres formations of Asturias have yielded a rich vertebrate fauna, represented by both abundant tracks and osteological remains. However, skeletal remains of theropod dinosaurs are rare, and the diversity of theropod tracks has only partially been documented in the literature. Here we describe the only non-dental osteological theropod remain recovered so far, an isolated anterior caudal vertebra, as well as the largest theropod tracks found. The caudal vertebra can be shown to represent a megalosaurine megalosaurid and represents the largest theropod skeletal remain described from Europe so far. The tracks are also amongst the largest theropod footprints reported from any setting and can be assigned to two different morphotypes, one being characterized by its robustness and a weak mesaxony, and the other characterized by a strong mesaxony, representing a more gracile trackmaker. We discuss the recently proposed distinction between robust and gracile large to giant theropod tracks and their possible trackmakers during the Late Jurassic-Berriasian. In the absence of complete pedal skeletons of most basal tetanurans, the identity of the maker of Jurassic giant theropod tracks is difficult to establish. However, the notable robustness of megalosaurine megalosaurids fits well with the described robust morphotypes, whereas more slender large theropod tracks might have been made by a variety of basal tetanurans, including allosaurids, metriocanthosaurids or afrovenatorine megalosaurids, or even exceptionally large ceratosaurs. Concerning osteological remains of large theropods from the Late Jurassic of Europe, megalosaurids seem to be more abundant than previously recognized and occur in basically all Jurassic deposits where theropod remains have been found, whereas allosauroids seem to be represented by allosaurids in Western Europe and metriacanthosaurids in more eastern areas. Short-term fluctuations in sea level might have allowed exchange of large theropods between the islands that constituted Europe during the Late Jurassic.

A Theropod Dinosaur from the Late Jurassic Cañadón Calcáreo Formation of Central Patagonia, and the Evolution of the Theropod Tarsus

Abstract. A fragmentary postcranial skeleton from the Late Jurassic (Oxfordian—Tithonian) Canadon Calcareo Formation of Chubut, Argentina, represents a new taxon of theropod dinosaur, which is here described as Pandoravenator fernandezorum gen. et sp. nov. This material represents the first Late Jurassic theropod known from Argentina. Pandoravenator fernandezorum is characterized by strongly elongated postzygapophyses in the caudal vertebrae and an unusual tarsal joint, with the astragalus showing two distal tubercles and a very low and laterally inclined ascending process. Phylogenetic analyses indicate that the new taxon is a basal tetanuran, although its exact phylogenetic position within basal tetanurans remains uncertain, due to the fragmentary nature of the remains and the lack of consensus among the different phylogenetic analyses. The tarsus of P. fernandezorum shows an intermediate morphology between that of basal theropods and more derived tetanurans. It is especially noteworthy for the presence...

Isolated theropod teeth associated with sauropod remains from El Oterillo II (Early Cretaceous) site of Salas de los Infantes (Burgos, Spain)

The relationship between dinosaur carcasses and isolated theropod teeth is well known in the fossil record. The usual explanation is that theropod dinosaurs fed on the herbivore carcass. Other evidence is provided by theropod tooth marks on the surface of herbivore skeletal remains. In this work we study isolated theropod teeth and the tooth marks in evidence on the bones of the sauropod from El Oterillo II (Salas de los Infantes, Spain, Lower Cretaceous). Theropod and crocodylomorph teeth have been found in relationship with the sauropod carcass. The fossils lie on channel lag deposits composed of sandstones with quartzite gravel. Thirty theropod teeth have been studied using qualitative features, statistical and cladistics analysis in addition to the tooth marks present on the vertebrae. The morphology of the theropod teeth has revealed greater palaeobiodiversity in these faunas than previously known, including baryonychine spinosaurids, basal tetanurans, dromaeosaurids and a singular coelurosaurian. The presence of tooth marks and isolated theropod teeth in close relationship with the sauropod carcass could also provide new evidence of the scavenging of theropod dinosaurs on the sauropods of this age and location. Six morphotypes of theropod teeth have been distinguished; the combination of basal and derived tetanurans is congruent with the known record from the Early Cretaceous of the Iberian Peninsula. The most probable explanation for the association of isolated theropod teeth and sauropod remains is the scavenging of the carcass by theropod dinosaurs.

A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the "middle" Cretaceous of Morocco.

Sigilmassasaurus brevicollis is an enigmatic theropod dinosaur from the early Late Cretaceous (Cenomanian) of Morocco, originally based on a few isolated cervical vertebrae. Ever since its original description, both its taxonomic validity and systematic affinities were contentious. Originally considered to represent its own family, Sigilmassasauridae, the genus has variously been suggested to represent a carcharodontosaurid, an ornithischian, and, more recently, a spinosaurid. Here we describe new remains referrable to this taxon and re-evaluate its taxonomic status and systematic affinities. Based on the new remains, a re-evaluation of the original materials, and comparisons with other spinosaurids, the holotype of Sigilmassasaurus brevicollis is identified as an anterior dorsal, rather than a cervical vertebra, and differences between elements referred to this taxon can be explained by different positions of the elements in question within the vertebral column. Many characters used previously to diagnose the genus and species are found to be more widespread among basal tetanurans, and specifically spinosaurids. However, the taxon shows several autapomorphies that support its validity, including the presence of a strongly rugose, ventrally offset triangular platform that is confluent with a ventral keel anteriorly in the mid-cervical vertebral centra and a strongly reduced lateral neural arch lamination, with no or an incomplete distinction between anterior and posterior centrodiapophyseal laminae in the posterior cervical and anterior dorsal vertebrae. We argue furthermore that Spinosaurus maroccanus, also described on the basis of isolated cervical vertebrae from the same stratigraphic unit and in the same paper as Sigilmassasaurus brevicollis, is a subjective synonym of the latter. Both a detailed comparison of this taxon with other theropods and a formal phylogenetic analysis support spinosaurid affintities for Sigilmassasaurus. However, we reject the recently proposed synonymy of both Spinosaurus maroccanus and Sigilmassasurus brevicollis with Spinosaurus aegyptiacus from the Cenomanian of Egypt, as there are clear differences between the vertebrae of these taxa, and they do not share any derived character that is not found in other spinosaurids. Together with a comparison with other spinosaurid vertebral material from the Kem Kem, this suggests that more than one taxon of spinosaurid was present in the Kem Kem assemblage of Morocco, so the referral of non-overlapping material from this unit to a single taxon should be regarded with caution.

An enigmatic plant-eating theropod from the Late Jurassic period of Chile.

Theropod dinosaurs were the dominant predators in most Mesozoic era terrestrial ecosystems. Early theropod evolution is currently interpreted as the diversification of various carnivorous and cursorial taxa, whereas the acquisition of herbivorism, together with the secondary loss of cursorial adaptations, occurred much later among advanced coelurosaurian theropods. A new, bizarre herbivorous basal tetanuran from the Upper Jurassic of Chile challenges this conception. The new dinosaur was discovered at Aysén, a fossil locality in the Upper Jurassic Toqui Formation of southern Chile (General Carrera Lake). The site yielded abundant and exquisitely preserved three-dimensional skeletons of small archosaurs. Several articulated individuals of Chilesaurus at different ontogenetic stages have been collected, as well as less abundant basal crocodyliforms, and fragmentary remains of sauropod dinosaurs (diplodocids and titanosaurians).

New theropod remains from the Tiourarén Formation (?Middle Jurassic, Niger) and their bearing on the dental evolution in basal tetanurans

A fragment of a maxilla and isolated theropod teeth from the (?) Middle Jurassic Tiourarén Formation are described. The specimens come from Tadibene, in the rural community of Aderbissinat, Thirozerine Department, Agadez Region, Niger. They were identified through direct comparison with teeth previously described in the literature as well as on the basis of discriminant and morphometric analyses. Our results suggest they belong to Ceratosauridae, Megalosauridae, and the oldest representatives of Spinosauridae. The analyzed sample shows some uncommon characters, such as spinosaurid-like ornamentation in megalosaurid-like teeth, or spinosaurid-like teeth with a low number of denticles, which sheds light on tooth morphology and dental evolution in basal tetanurans and early spinosaurids.

Large sauropod and theropod tracks from the Middle Jurassic Chuanjie Formation of Lufeng County, Yunnan Province and palaeobiogeography of the Middle Jurassic sauropod tracks from southwestern China

Tracks of large theropods and a single sauropod footprint are reported from red beds at Beikeshan locality in the Middle Jurassic Chuanjie Formation, of Lufeng County, near the large World Dinosaur Valley Park complex. The Chuanjie theropod tracks are assigned to the ichnogenus Eubrontes and the large sauropod track is given the provisional label Brontopodus. All occur as isolated tracks, i.e., trackways are not preserved. Saurischian dominated ichnofaunas are relatively common in the Jurassic of China. The producers of the Chuanjie tracks may have been similar to the basal tetanuran theropod Shidaisaurus and to mamenchisaurid sauropods, which were widely distributed throughout China, during the Jurassic, and are known from skeletal remains found in the same unit. Other potential sauropod trackmakers include titanosauriforms or as-yet-unknown basal eusauropods. The ichno- and skeletal records from the Jurassic of the Lufeng Basin are largely consistent, and both document the presence of middle-large sized theropods and sauropods.

The phylogeny of Tetanurae (Dinosauria: Theropoda)

Tetanuran theropods represent the majority of Mesozoic predatory dinosaur diversity and the lineage leading to extant Aves. Thus their history is relevant to understanding the evolution of dinosaur diversity, Mesozoic terrestrial ecosystems, and modern birds. Previously, the fragmentary and poorly sampled fossil record of basal (non-coelurosaur) tetanurans led to uncertainties regarding their basic interrelationships. This in turn prevented determining the relationships of many incompletely known taxa that nonetheless document a global radiation spanning more than 120 million years. We undertook an exhaustive examination of all basal tetanurans and all existing character data, taking advantage of recent discoveries and adding new morphological, temporal and geographic data. Our cladistic analysis of 61 taxa achieved significantly improved phylogenetic resolution. These results position several ‘stem’ taxa basal to a succession of monophyletic clades (Megalosauroidea, Allosauroidea and Coelurosauria). Megalosauroids include nearly 20 taxa arrayed amongst a basalmost clade (Piatnitzkysauridae, fam. nov.) and the sister taxa Spinosauridae and Megalosauridae; the latter includes two subfamilies, Megalosaurinae and Afrovenatorinae subfam. nov. Allosauroidea contains a diverse Metriacanthosauridae (= Sinraptoridae), Neovenatoridae, Carcharodontosauridae and a reduced Allosauridae. Finally, we assessed more than 40 fragmentary forms and hundreds of additional reported tetanuran occurrences. Tetanuran evolution was characterized by repeated acquisitions of giant body size and at least two general skull forms, but few variations in locomotor morphology. Despite parallel diversification of multiple lineages, there is evidence for a succession of ‘dominant’ clades. Tetanurae first appeared by the Early Jurassic and was globally distributed by the Middle Jurassic. Several major clades appeared prior to the breakup of Pangaea; as such their absence in specific regions, and at later times, must be due to poor sampling, dispersal failure and/or regional extinction. Finally, we outline a general perspective on Mesozoic terrestrial biogeography that should apply to most clades that appeared before the Late Jurassic.

The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the early cretaceous of Laos

Spinosaurids are among the largest and most specialized carnivorous dinosaurs. The morphology of their crocodile-like skull, stomach contents, and oxygen isotopic composition of the bones suggest they had a predominantly piscivorous diet. Even if close relationships between spinosaurids and Middle Jurassic megalosaurs seem well established, very little is known about the transition from a generalized large basal tetanuran to the specialized morphology of spinosaurids. Spinosaurid remains were previously known from the Early to Late Cretaceous of North Africa, Europe, and South America. Here, we report the discovery of a new spinosaurid theropod from the late Early Cretaceous Savannakhet Basin in Laos, which is distinguished by an autapomorphic sinusoidal dorsosacral sail. This new taxon, Ichthyovenator laosensis gen. et sp. nov., includes well-preserved and partially articulated postcranial remains. Although possible spinosaurid teeth have been reported from various Early Cretaceous localities in Asia, the new taxon I. laosensis is the first definite record of Spinosauridae from Asia. Cladistic analysis identifies Ichthyovenator as a member of the sub-clade Baryonychinae and suggests a widespread distribution of this clade at the end of the Early Cretaceous. Chilantaisaurus tashouikensis from the Cretaceous of Inner Mongolia, and an ungual phalanx from the Upper Jurassic of Colorado are also referred to spinosaurids, extending both the stratigraphical and geographical range of this clade.

A Reassessment ofKelmayisaurus petrolicus, a Large Theropod Dinosaur from the Early Cretaceous of China

The Early Cretaceous fossil record of large-bodied theropods from Asia is poor, hindering comparison of Asian predatory dinosaur faunas with those from other continents. One of the few large Asian theropod specimens from this interval is a partial skull (maxilla and dentary) from the Lianmugin Formation (?Valanginian—Albian), the holotype of Kelmayisaurus petrolicus. Most authors have either considered this specimen as an indeterminate basal tetanuran or a nomen dubium. We redescribe K. petrolicus and note that it possesses a single autapomorphy (a deep accessory groove on the lateral surface of the anterior dentary), as well as a unique combination of characters that differentiates it from other theropods, affirming its validity. A phylogenetic analysis recovers K. petrolicus as a basal carcharodontosaurid, which is supported by various features: very deep interdental plates (a carcharodontosaurid synapomorphy), fused interdental plates (present in carchardontosaurids and a limited number of other therop...

An abelisauroid dinosaur from the Middle Jurassic of Laurasia and its implications on theropod palaeobiogeography and evolution

We review here the phylogenetic relationships of a theropod distal end of tibia (MB. R.2351) from the Middle Jurassic Stonesfield Slate (Taunton Formation, middle Bathonian) of Oxfordshire, England. This specimen was previously described as a small basal tetanuran, but our reinterpretation suggests that it was an early member of the Abelisauroidea. The new assignment is supported by the presence of an apomorphic vertical facet for the reception of the ascending process of the astragalus, sub-rectangular anterior scar of the astragalar ascending process, median vertical ridge in the scar for the reception of the ascending process of the astragalus, and posterolateral process not distinctly offset from the lateral margin of the shaft. In particular, the Stonesfield specimen shares an overall morphology and a unique combination of apomorphies with the Middle Jurassic Australian abelisauroid Ozraptor. Nevertheless, both specimens differ in some punctual features. MB. R.2351 constitutes the oldest evidence of an abelisauroid outside Gondwana and indicates that the group had achieved a Pangean distribution during, at least, the Middle Jurassic. Thus, the initial diversification of abelisauroids would have occurred earlier than previously thought. Accordingly, the Middle and Late Jurassic Pangean distribution of abelisauroids implies that the absence of the group in Cretaceous Asiamerican assemblages would reflect a regional extinction, in which a competitive replacement with coelurosaurs (e.g. tyrannosauroids) is surely one of the hypotheses that should be tested in future studies.

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.