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.     

The oldest theropods from the Neuquén Basin: Predatory dinosaur diversity from the Bajada Colorada Formation (Lower Cretaceous: Berriasian–Valanginian), Neuquén,Argentina

Here, we describe new theropod materials (several isolated teeth, an axis, two caudal centra and a proximal left tibia) from the type locality of the Bajada Colorada Formation (Lower Cretaceous, Berriasian–Valanginian). Although fragmentary, the recovered material shows a diverse association of meat-eating dinosaurs for this poorly understood period of time. Three techniques were used to assess the phylogenetic position of the isolated teeth: multivariate (PCA), discriminant and phylogenetic analyses. The morphotypes 1, 2 and 3 (small non-recurved isolated crowns) were regarded as Theropoda indet., as our analyses failed to support a more precise classification. Two large almost complete ziphodont crowns, considered morphotype 4, were identified as belonging to megalosaurid tetanurans by phylogenetic, discriminant and multivariate analyses, thus likely representing the first record of this ancient family in South America, and the youngest worldwide. We refer the axis to a small abelisauroid ceratosaurian based on the following suite of characters; long and pointed epipophyses, a pneumatic foramen in the centrum, the invaginated spinopostzygapophyseal lamina, and the anteroposteriorly long, dorsally convex, and dorsally/posteriorly unexpanded neural spine. The caudal centra can be referred to a theropod, and tentatively to Abelisauroidea, suggesting that they may belong to the same taxon than that represented by the axis. The tibia lacks the incisura tibialis, presents a low, sharp and proximally positioned fibular crest, and has a deep lateral fossa, which is limited dorsally by a coarse, rounded in section and anteriorly directed crest. This combination of features allows to refer it to a large abelisaurid. The Bajada Colorada dinosaur record includes so far a small abelisauroid, a large abelisaurid, a probably medium to large megalosaurid tetanuran, diplodocid and dicraeosaurid sauropods. It shows some similarities with Middle and Upper Jurassic units in central Patagonia, Africa and Portugal, suggesting that no significant dinosaur faunal turnover took place through the Jurassic-Cretaceous event in the southern part of South America.     

First Brazilian carcharodontosaurid and other new theropod dinosaur fossils from the Campanian–Maastrichtian Presidente Prudente Formation,São Paulo State,southeastern Brazil

We report new theropod dinosaur material from the Presidente Prudente Formation (Campanian–Maastrichtian), Bauru Group, in southwestern São Paulo state. The material comprises a fragment of right maxilla of Carcharodontosauridae, an anterior portion of a left ilium of Abelisauroidea and a proximal portion of a right fibula of a coelurosaurian. Previous theropod records from the Bauru Basin comprise Abelisauridae and Tetanurae and in São Paulo state these have been represented by only a right premaxilla of an abelisaurid and isolated abelisaurid and carcharodontosaurid teeth. The new material reported here represents the first theropod remains from the Presidente Prudente Formation, and includes the first abelisauroid and coelurosaurian postcranial remains from the Bauru Basin in São Paulo state and the first latest Cretaceous carcharodontosaurid known from non-dental remains anywhere.     

First elaphrosaurine theropod dinosaur (Ceratosauria: Noasauridae) from Australia — A cervical vertebra from the Early Cretaceous of Victoria

Elaphrosaurinae is an enigmatic clade of gracile ceratosaurian theropod dinosaurs known from the Late Jurassic of Africa (Elaphrosaurus bambergi) and Asia (e.g., Limusaurus inextricabilis), and the early Late Cretaceous of Argentina (Huinculsaurus montesi). Elaphrosaurinae is often placed within Noasauridae as the sister taxon to Noasaurinae, a clade of small-bodied theropods that lived in South America, Africa, Madagascar and India throughout much of the Cretaceous. Herein, we report the first evidence of Elaphrosaurinae from Australia: a nearly complete middle cervical vertebra from the upper Lower Cretaceous (lower Albian) Eumeralla Formation of Cape Otway, Victoria, Australia. The fact that this site would have been situated at ~76°S towards the end of the Early Cretaceous (~110–107 Ma) implies that elaphrosaurines were capable of tolerating near-polar palaeoenvironments, whereas its age indicates that elaphrosaurines persisted in Australia until at least the late Early Cretaceous. The new Australian elaphrosaurine, in tandem with the recently described Huinculsaurus montesi from the Cenomanian–Turonian of Argentina, implies that the spatiotemporal distribution of Elaphrosaurinae has heretofore been greatly underestimated. Historic confusion of elaphrosaurines with coelurosaurs, especially ornithomimosaurs, coupled with our generally poor understanding of noasaurid evolution, might explain the apparent dearth of fossils of this theropod clade worldwide.     

Ornithomimidae (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan

The stratigraphically oldest remains of ornithomimid theropod dinosaurs are known from the Cenomanian Khodzhakul Formation and the Turonian Bissekty Formation of Uzbekistan. The ornithomimid from the Bissekty Formation is documented by more than 800 isolated bones that represent much of the skeleton. It shows at least three unambiguous synapomorphies of Ornithomimidae: length of anterior cervical centra three to five times greater than transverse width; low and rounded fibular crest of tibia; metatarsal III pinched between metatarsals II and IV; and proximal end of metatarsal III not visible in anterior view. Phylogenetic analysis, based on a dataset with 568 morphological characters and including all known ornithomimosaurian taxa, places the ornithomimid from the Bissekty Formation near the base of the ornithomimid radiation, between Archaeornithomimus asiaticus and Sinornithomimus dongi.     

The oldest known metriorhynchid crocodylian from the Middle Jurassic of North-eastern Italy: Neptunidraco ammoniticus gen. et sp. nov.

Metriorhynchidae is a clade of marine-adapted crocodilians known from several Middle Jurassic–Early Cretaceous specimens collected predominantly in South America and Europe, but poorly known in the northern margin of Gondwana. The “Portomaggiore crocodile” is the most complete specimen of an Italian metriorhynchid to date: it consists of a partial skeleton that has been provisionally referred to an unnamed species of Late Jurassic Metriorhynchus or Geosaurus. The specimen is preserved in the reddish, nodular limestone of the Rosso Ammonitico Veronese Formation (Bajocian–Tithonian); new data on microfossil associations constrain the age of the metriorhynchid to the late Bajocian–earliest Bathonian. On the basis of cranial synapomorphies, the “Portomaggiore crocodile” falls as the closest sister-taxon of the Late Jurassic–Early Cretaceous geosaurines, and is referred to Neptunidraco ammoniticus gen. et sp. nov. It is unique among Middle Jurassic metriorhynchids in showing an incipient streamlining of the skull, shared with Late Jurassic and Cretaceous taxa. Since Neptunidraco is the oldest known member of Metriorhynchidae, its phylogenetic position supports the hypothesis that the timing of the initial metriorhynchid and geosaurine diversifications should start in the Bajocian.     

A diversified vertebrate ichnite fauna from the Feitianshan Formation (Lower Cretaceous) of southwestern Sichuan,China

The Bajiu tracksite exposed on a fine sandstone surface at the top of the Feitianshan Formation, close the contact with the overlying Xiaoba Formation, reveals a saurischian dominated ichnofauna consisting of four sauropod and at least eight theropod trackways. The sauropod trackways display a medium-wide gauge pattern characteristic of titanosauriforms, while the theropod trackways can be subdivided into two distinct morphotypes. One is similar to the ichnogenera Eubrontes and Megalosauripus, the other (cf. Dromaeopodus) appears to represent a functionally didactyl dromaeosaur where pedal digit II is represented only by an oval basal pad. A single swim track is possibly attributable to a turtle or crocodylian. Such sauropod-theropod dominated ichnofaunas are consistently typical of the red-bed tracksites in the region and useful for characterizing a fauna that is otherwise poorly known from body fossils. The stratigraphic position and late occurrence of the (typical Jurassic) Eubrontes-Megalosauripus morphotype is well known from other Cretaceous localities in China and seems to reflect a peculiarity in the theropod faunas of East Asia.     

Troodontidae (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan

To date three taxa of troodontid theropod dinosaurs have been recognized from Upper Cretaceous strata in two regions of the Kyzylkum Desert in Uzbekistan. The Cenomanian Khodzhakul Formation in the southwestern Kyzylkum Desert has yielded isolated serrated teeth and some postcranial bones of an indeterminate troodontid. In the central Kyzylkum Desert troodontids are known from the Cenomanian Dzharakuduk Formation (Urbacodon itemirensis) and the Turonian Bissekty Formation (Urbacodon sp.). Urbacodon itemirensis is known from a single dentary whereas Urbacodon sp. is represented by isolated teeth, maxilla and dentary fragments, a partial braincase, and some postcranial bones. The troodontid affinities of Urbacodon are supported by several synapomorphies: presence of a subotic recess; reduced basal tubera placed directly under the occipital condyle; maxilla participating in the margin of the external naris; nutrient foramina on dentary situated within a deep lateral groove; dentary without distinct interdental plates; large number of small dentary and maxillary teeth; teeth constricted between root and crown; anterior dentary teeth smaller, more numerous, more closely spaced than those in the middle of the tooth row, and implanted in a groove; posterior dorsal vertebrae with tall and posterodorsally tapering neural spines; and presence of a midline sulcus on the neural arches of distal caudals. Among Troodontidae, Urbacodon resembles Byronosaurus, Gobivenator, and Xixiasaurus in the absence of serrations on the tooth crowns and having premaxillary teeth that are D-shaped in cross-section. However, phylogenetic analysis did not recover a clade of Asiatic troodontids with unserrated teeth.     

Morphology and internal structure of two new abelisaurid remains (Theropoda,Dinosauria) from the Adamantina Formation (Turonian – Maastrichtian), Bauru Group,Paraná Basin,Brazil

The knowledge on the Brazilian theropod fauna is hampered by the limited number of specimens unearthed so far. The most potential deposits for the finding of those dinosaurs are the layers of Bauru Group, which comprises several different formations of Late Cretaceous age. Most of those remains are referred to Abelisauroidea, a clade that is particularly well represented in Gondwana. Here we report two new abelisaurid specimens comprising a left ilium (DGM 927-R) and the distal articulation of a right femur (MCT 1857-R) unearthed from the outcrops of Adamantina Formation (Turonian – Maastrichtian) in the locality known as Santo Anastácio, that comprises an abandoned quarry located in the outskirts of the homonymous city in São Paulo State. The ilium has a preacetabular process horizontally oriented and ventrally deflected, as well as an acute angle between this process and the pubic peduncle. The femur shows a well-developed anteromedial crest, but not to the same degree as in noasaurids, and ossified ridges between condyles and tibiofibular crest. Those features allow us to assign both specimens to Abelisauridae. CT-scans of those specimens shows that the ilium has pneumaticities similar to the camellate pattern previously recorded in neosauropods and reveals the diverticula invasion of abdominal air-sacs. The femur presents a developed anisotropic trabecular system, which suggests that the main muscular strain is located at the distal portion of this bone. These features have not been previously reported in Abelisauridae, indicating that the internal organization of the postcranial skeleton in the group is rather complex and quite variable.     

Theropod dinosaur diversity and palaeobiology in the Wealden Group (Early Cretaceous) of England: evidence from a previously undescribed tibia

A robust, partial right tibia of a theropod dinosaur (Natural History Museum London collections, BMNH R9385) is described for the first time. The specimen was collected at Hastings, Sussex (England) in the last century, and is among the oldest known of English Wealden Group theropods. It represents a tetanuran theropod that may have been about 3 m in total length, and is distinct from all currently known Wealden theropods for which tibiae have been described. The present specimen is significant palaeobiologically in exhibiting a series of theropod tooth marks on its caudal surface, indicating predation or scavenging by another theropod.     

The Late Jurassic ray Kimmerobatis etchesi gen. et sp. nov. and the Jurassic radiation of the Batoidea

The laminated marine mudstones of the Late Jurassic of Kimmeridge, southern England, yield two exceptionally well-preserved partial skeletons of a previously unrecognised species of early batoid. These are described as a new genus and species, Kimmerobatis etchesi gen. et sp. nov. which has a general “guitarfish” bauplan as in all other batoids known from the Jurassic. This species possesses a combination of primitive characters such as centra present within the majority of the synarcual and antorbital cartilages that fail to reach the pectoral skeleton along with more derived characters, such as the lack of fin spines. Until now, little study has been carried out on the affinities of Jurassic batoids, despite their key role in understanding batoid evolution. Results from parsimony and likelihood phylogenetic reconstruction indicates that the whole-bodied Jurassic batoids Spathobatis, Belemnobatis, and Kimmerobatis gen. nov. form their own clade, Spathobatidae, and do not lend support to a monophyletic “Rhinobatidae”. Among Jurassic batoids, Kimmerobatis gen. nov. is most derived, but with derived characters being independently acquired compared to modern batoids (e.g. presence of a postpelvic process). The inclusion of whole bodied Jurassic fossils have generated a more resolved hypothesis of batoid evolution throughout the Cretaceous and into the Cenozoic.     

The longest theropod trackway from East Asia,and a diverse sauropod-, theropod-, and ornithopod-track assemblage from the Lower Cretaceous Jiaguan Formation,southwest China

Here we report a large dinosaur tracksite from an extensive fluvial sandstone surface in the Lower Cretaceous Jiaguan Formation of Sichuan Province, China. The site contains over 250 individual tracks comprising at least 18 recognizable trackways, including the longest theropod trackway (cf. Eubrontes) known from China. This exceptional theropod trackway consists of 81 successive footprints covering a distance of 69 m. The tracks are well-preserved and are expressed both as true tracks on the main “upper” surface and as transmitted undertracks on a locally exposed “lower” bed. Also recorded are six other theropod trackways, including small Grallator-like ichnites, eight sauropod trackways (cf. Brontopodus), and three small ornithopod (cf. Ornithopodichnus) trackways with a parallel orientation, which may indicate gregarious behavior. Several trackways of a larger theropod trackmaker show pes imprints with elongated traces of the metatarsals, suggesting extramorphological (substrate-controlled) variation and/or plantigrade posture, which is here interpreted as indicating a change in gait assumed in response to deep and soft sediment. The assemblage indicates a diverse dinosaur fauna in the Lower Cretaceous Sichuan Basin with variously sized theropods, sauropods, and ornithopods. The late occurrence of footprints of the Grallator-Eubrontes plexus in Lower Cretaceous strata is further evidence of the extended stratigraphic range of this morphotype and the distinct palaeobiogeographic distribution of these trackmakers in East Asia.     

New rebbachisaurid (Dinosauria: Sauropoda) material from the Wessex Formation (Barremian,Early Cretaceous), Isle of Wight,United Kingdom

Rebbachisauridae is a poorly understood clade of diplodocoid sauropod dinosaurs, currently known only from the Cretaceous of Africa, Europe and South America. European representatives are particularly rare and fragmentary. Here, we report an anterior caudal vertebra from the Barremian (Early Cretaceous) Wessex Formation of the Isle of Wight, off the southern coast of England. This specimen possesses several features known only in rebbachisaurids and shares two synapomorphies with the Afro-European taxa Demandasaurus darwini and Nigersaurus taqueti, both pertaining to the morphology of the neural spine. These features are the development of triangular lateral processes and the presence of an elliptical fossa on the lateral surface, bounded by the lateral lamina and postspinal rugosity. The Isle of Wight specimen also shares several features solely with Demandasaurus, indicating a close relationship with the Spanish taxon. These include the presence of a hyposphenal ridge, as well as an anteriorly excavated caudal rib that is restricted almost entirely to the neural arch. However, it differs from Demandasaurus in a number of ways, including the lack of excavation on the posterior surface of the caudal rib, the orientation of the neural spine, and the composition and morphology of the lateral lamina. In addition, the Isle of Wight vertebra possesses one potential autapomorphy: bifurcation of the elliptical fossa on the neural spine. However, because of the fragmentary nature of the material, a new name is not erected. Along with Demandasaurus and Histriasaurus boscarollii, this caudal vertebra indicates the presence of at least three European rebbachisaurid taxa and provides new anatomical information on this enigmatic clade of sauropod dinosaurs.     

A new, large tyrannosaurine theropod from the Upper Cretaceous of China

Tyrannosaurids are primarily gigantic, predatory theropod dinosaurs of the Cretaceous. Here we report a new member of the tyrannosaurid clade Tyrannosaurinae from the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China, based on a maxilla and associated dentary. The discovery of this animal, here named Zhuchengtyrannus magnus gen. et sp. nov., adds to the known diversity of tyrannosaurids in Asia. Z. magnus can be identified by a horizontal shelf on the lateral surface of the base of the ascending process, and a rounded notch in the anterior margin of the maxillary fenestra. Several additional features contribute to a unique combination of character states that serves to further distinguish Z. magnus from other taxa. Comparisons with other tyrannosaurids suggest that Zhuchengtyrannus was a very large theropod, comparable in size to both Tarbosaurus and Tyrannosaurus.     

A large-clawed theropod (Dinosauria: Tetanurae) from the Lower Cretaceous of Australia and the Gondwanan origin of megaraptorid theropods

Megaraptoridae comprises a clade of enigmatic Gondwanan theropods with characteristic hypertrophied claws on the first and second manual digits. The majority of megaraptorids are known from South America, although a single genus (Australovenator) plus additional indeterminate material is also known from Australia. This clade has a controversial placement among theropods, and recently has been interpreted alternatively as a carcharodontosaurian or a tyrannosauroid lineage. We describe new fragmentary but associated postcranial remains from the opal fields of Lightning Ridge (middle-Albian, Griman Creek Formation) in north-central New South Wales. The new unnamed taxon exhibits a number of unusual features that suggest the presence of a hitherto unrecognised Australian megaraptorid. From an Australian perspective, the Lightning Ridge taxon predates Australovenator by ca. 10 Ma and is minimally coeval with megaraptoran material reported from the Eumeralla Formation of Victoria (but potentially 6.1–9.5 Ma younger). It is also notable as the largest predatory dinosaur yet identified from Australia and is only the second theropod known from more than a single element. A Bayesian phylogenetic approach integrating morphological, stratigraphic and palaeogeographic information tested both the carcharodontosaurian and tyrannosauroid placements for Megaraptora. Regardless of the preferred placement among Tetanurae, rigorous palaeobiogeographic analyses support an Asian origin of Megaraptora in the latest Jurassic (about 150–135 Ma), an Early Cretaceous (about 130–121 Ma) divergence of the Gondwanan lineage leading to Megaraptoridae, and an Australian root for megaraptorid radiation. These results indicate that Australia's Cretaceous dinosaur fauna did not comprise simply of immigrant taxa but was a source for complex two-way interchange between Australia–Antarctica–South America leading to the evolution of at least one group of apex predatory dinosaurs in Gondwana.     

A new hangingfly (Insecta,Mecoptera, Bittacidae) from the Purbeck Limestone Group (Lower Cretaceous) of southern England and a review of Cretaceous Bittacidae

Rapid radiation of the Bittacidae during the Jurassic resulted in high diversity within this family of Mecoptera. More than 40 species within 23 genera have been described from this period. However, around the end of the Jurassic the abundance and diversity of Bittacidae decreased and only seven species in five genera are known to date from the Cretaceous. A new specimen from the basal Cretaceous of England, Tytthobittacus jarzembowski sp. nov., described here, represents the eighth species and the second fossil representative of this family from the European Cretaceous. This hangingfly belongs to an extant and relictual genus previously known only from Australia. The paper also includes a review of all known Cretaceous bittacids and a re-examination of European species Antiquanabittacus nanus Petrulevičius and Jarzembowski, 2004.     

A hypertrophied ungual phalanx from the lower Barremian of Spain: Implications for the diversity and palaeoecology of Spinosauridae (Theropoda) in Iberia

An enlarged theropod manual ungual (CSC1-4) from the Weald facies of Spain is described. The claw was found in the fossil locality of Caña Seca 1, Teruel province, within the El Castellar Formation of early Barremian (Early Cretaceous) in age.CSC1-4 is morphologically closer to megalosauroids than to any other theropod clade bearing enlarged manual claws and shows the greatest similarity to the manual ungual of digit I of Baryonyx walkeri. Both CS1-4 and this taxon share a particularly enlarged, elongated and transversely wide manual claw. CSC1-4, however, differs from Baryonyx's ungual in having less curvature, a straight dorsal edge in the proximal part, slightly more width above the grooves than below, and a certain asymmetry, with the lateral face more flattened. Taking into account the palaeogeographic and temporal context, these considerations suggest that they are closely related but distinct spinosaurid taxa.The presence of an enlarged manual claw in spinosaurids has been invoked as an anatomical feature typically associated with scavenging and hunting habits, as well as digging behaviour. The spinosaurid record from the Barremian of the Iberian Peninsula shows that members of this clade favored freshwater environments with some marine influence in this part of Europe.     

A New Theropod Dinosaur from the Middle Jurassic of Lufeng,Yunnan, China

A new theropod dinosaur, Shidaisaurus jinae gen. et sp. nov., has been described on the basis of an incomplete skeleton. The specimen was found near the base of the Upper Lufeng Formation (early Middle Jurassic) in Yunnan, China. It is the first theropod dinosaur from the Middle Jurassic of Yunnan. Shidaisaurus jinae is distinguishable from other Jurassic theropods by certain features from the braincase, axis, and pelvic girdle. The absence of any pleurocoels in the axis or in any anterior dorsal vertebrae suggests that the new Lufeng theropod is relatively primitive and more plesiomorphic than most of the Middle to Late Jurassic theropods from China. Most Chinese taxa of Jurassic theropod dinosaurs have not been well described; a further detailed study will be necessary for us to determine their phylogenetic relationships with Shidaisaurus jinae.     

A tooth of Archaeopterygidae (Aves) from the Lower Cretaceous of France extends the spatial and temporal occurrence of the earliest birds

The Late Jurassic Archaeopterygidae, comprising the iconic genus Archaeopteryx, is altogether among the earliest, basalmost and best-known Mesozoic avian taxa. The geographic distribution of Archaeopteryx is hitherto restricted to a limited region of southern Germany, probably due to preservation biases. Here we describe a tooth sorted among the more than 35,000 isolated teeth found by sieving sediments from the Cherves-de-Cognac locality (western France, Lower Cretaceous). This tooth crown is morphologically similar to teeth of the German specimens of Archaeopteryx (sensu lato), despite minor differences. The Cherves-de-Cognac tooth differs much more importantly from all other known taxa. It shares with different specimens of Archaeopteryx the general tooth size, recurved shape affecting apical third of crown, thin apical-mesial carina, constriction at base of crown giving distinctive S shape of distal edge in profile, absence of other ornamentation or serration, and shape of crown section. Incidentally, former assignments of teeth from the Upper Jurassic of Guimarota (Portugal) to cf. Archaeopteryx are unwarranted, as those teeth markedly differ from the Archaeopterygidae in several crucial features. We assign the new tooth to the family Archaeopterygidae, the earliest European birds, making it the first member of the family in Europe outside Germany, and extending its temporal occurrence to the early Cretaceous.