A protoceratopsid skeleton with an associated track from the Upper Cretaceous of Mongolia

The Djadokhta Formation of the Gobi Desert is known for the number and diversity of dinosaur and other vertebrate bones and skeletons found there, but only theropod, hadrosaur and supposed ankylosaurid footprints have been reported from this stratum. Dinosaur footprints are also noted from the Nemegt Formation, and occur as typical dinosaur track accumulations (tracksites). An articulated protoceratopsid skeleton - specimen ZPAL Mg D-II/3 - was collected by the Polish-Mongolian Expedition of 1965 from the Djadokhta Formation of Flaming Cliffs in Mongolia. Recently, the natural cast of a tetradactyl digitigrade footprint was found underneath the pelvic girdle while the skeleton and matrix were being prepared. This is possibly the first find of a dinosaur track in close association with an articulated skeleton. Although Protoceratops is an extremely common dinosaur in Mongolia, its footprints have never previously been reported from the Late Cretaceous of the Gobi Desert.     

Juvenile specimens of Pinacosaurus grangeri Gilmore, 1933 (Ornithischia: Ankylosauria) from the Late Cretaceous of China, with comments on the specific taxonomy of Pinacosaurus

Four juvenile specimens referable to Pinacosaurus grangeri (Ankylosauria: Dinosauria) are described from the Campanian (Upper Cretaceous) locality Bayan Mandahu in northern Inner Mongolia Autonomous Region (People’s Republic of China). All the specimens preserve the skulls as well as, in some cases, mandibles, postcrania, and osteoderm. They are not taphonomically deformed by expanding matrix distortion, unlike many Gobi specimens, including the holotype of P. grangeri. Bayan Mandahu is also the type locality for Pinacosaurus mephistocephalus. The proximity in space and time of these two closely related species warrants a generic and specific revision for Pinacosaurus. The distinction of the two species is based on characters of the squamosal dermal elaborations, cranial roof posterior to the orbits, premaxillary notch, and distal margin of the ilium. Although a relatively well-represented ankylosaur taxon, the phylogenetic position of Pinacosaurus has not been unequivocally resolved. A new analysis recovers Pinacosaurus as the most basal member of the Ankylosaurinae.     

The cranial morphology and taxonomic status of Tarchia (Dinosauria: Ankylosauridae) from the Upper Cretaceous of Mongolia

The taxonomic status of specimens referred to Tarchia kielanae, T. gigantea, and Minotaurasaurus ramachandrani has been contested. The holotype of T. kielanae, Zaklad Paleobiologii (Institute of Paleobiology)-Polish Academy of Sciences (ZPAL) MgD I/111, is redescribed. It has features common to both Palaeontological Institute-Russian Academy of Sciences (PIN) 3142/250, long considered Tarchia, and to M. ramachandrani but which are lacking in Saichania chulsanensis. The specimen PIN 3142/250 is not referable to Saichania but instead represents a new species, Tarchia teresae sp. nov. Similarly, the holotype skull of M. ramachandrani is not referable to T. kielanae, so Minotaurasaurus is a valid taxon. Tarchia is more derived than either Saichania or Minotaurasaurus while sharing features of both.     

Reuniting the “head hunted” Corythosaurus excavatus (Dinosauria: Hadrosauridae) holotype skull with its dentary and postcranium

A Corythosaurus skull (UALVP 13) was collected in 1920 from what is now Dinosaur Provincial Park in southern Alberta, Canada and was designated the holotype of Corythosaurus excavatus, a new hadrosaur species by Gilmore (1923). In 1992, a previously uncovered, weathered, headless skeleton was found. This was collected in 2012 by the University of Alberta as there was potential it could belong to the holotype skull. In addition, an isolated hadrosaur dentary had been found in 1992 close to the articulated postcranial skeleton and may be one of the missing jaws of Corythosaurus excavatus. The hypothesis that it may be the skeleton of the holotype of Corythosaurus excavatus is tested using anatomical information and statistical analyses. Statistical comparisons suggest it is possible that the skull and dentary belong to the same specimen. Furthermore, bivariate plots and percentage prediction errors also indicate that the postcranial material could belong to the UALVP 13 skull. Because many large vertebrate fossil specimens (even types) are not always collected in their entirety, this method may be used as a line of evidence to determine whether independently collected specimens potentially belong to the same individual. The problems described herein highlight the need to collect specimens in their entirety, and for good field documentation, including the spatial and stratigraphic context of all finds.     

A New Species of the Ankylosaurid Dinosaur Crichtonsaurus(Ankyiosauridae:Ankylosauria)from the Cretaceous of Liaoning Province,China

A new ankylosaurid species of Crichtonsaurus,C.benxiensis sp.nov.,is erected based on a well preserved skeleton and a nearly complete skull found in the Sunjiawan Formation(Cenomanian- Turonian),Beipiao,Liaoning Province.Crichtonsaurus benxiensis sp.nov.is characterized by width 84.6% of length of the skull,a deep depression located medial to the notch between the orbit and the squamosal horn,paroccipital processes fused to the quadrates,fused scapula and coracoid,a large foramen for the supracoracoid nerve to enter the coracoid and to exit out from the scapula.Based on the new specimen,Crichtonsaurus is assigned to Ankylosauridae.     

A New Species of the Ankylosaurid Dinosaur Crichtonsaurus (Ankylosauridae: Ankylosauria) from the Cretaceous of Liaoning Province, China

A new ankylosaurid species of Crichtonsaurus, C. benxiensis sp. nov., is erected based on a well preserved skeleton and a nearly complete skull found in the Sunjiawan Formation (Cenomanian–Turonian), Beipiao, Liaoning Province. Crichtonsaurus benxiensis sp. nov. is characterized by width 84.6% of length of the skull, a deep depression located medial to the notch between the orbit and the squamosal horn, paroccipital processes fused to the quadrates, fused scapula and coracoid, a large foramen for the supracoracoid nerve to enter the coracoid and to exit out from the scapula. Based on the new specimen, Crichtonsaurus is assigned to Ankylosauridae.     

Osteology and ontogeny of Early Cretaceous Philydrosaurus (Diapsida: Choristodera) based on new specimens from Liaoning Province,China

Choristoderes are a group of extinct diapsid reptiles that once occupied the freshwater systems in the Northern Hemisphere from the Middle Jurassic through the Miocene. The Early Cretaceous monjurosuchid Philydrosaurus from western Liaoning, China, represents a transitional morphotype between a broad-snouted (crocodile-like) and a narrow-snouted (gavial-like) skull during the evolution of Choristodera. New specimens of the taxon from the Early Cretaceous Jiufotang Formation include an incomplete braincase and a nearly complete juvenile skeleton; the study of these specimens provides new information on the braincase and ontogenetic change of the skeleton of this transitional monjurosuchid. In the braincase floor, the foramen internus canalis caroticus externus opens in a groove lateral to the constricted lateral aspect of the parasphenoid, and ventral exposure of this foramen differs from Champsosaurus, in which parasphenoid/pterygoid fusion completely encloses the canal. On the occiput, the vagus nerve foramen penetrates the basioccipital, while the two foramina for the hypoglossal nerve open at the exoccipital/basioccipital suture. Comparative study of the new juvenile specimen with adult Philydrosaurus reveals previously unknown developmental changes of the cranial and postcranial skeleton of this monjurosuchid. Ontogenetically, Philydrosaurus underwent a dramatic change of skull proportions, including elongation of the antorbital and postorbital regions, and elongation of the jaws, with a great increase of the number of marginal teeth. The lower temporal fenestra is entirely closed in early ontogeny, as in large, fully-grown adults.     

A new ctenochasmatoid pterosaur from the Early Cretaceous Yixian Formation of western Liaoning,China

A new ctenochasmatoid pterosaur, Gladocephaloideus jingangshanensis gen. et sp. nov. from the Yixian Formation of western Laioning Province is erected based on a complete skull and partial postcranial skeleton. It is characterized by following features: about 50 total teeth with sharp tips; small nasoantorbital opening, occupying approximately 13% of the skull length; ratio of prenarial length to skull length approximately 0.63. The diagnoses of the Ctenochasmatoidea and Gallodactylidae are amended based on the new taxon. Gladocephaloideus jingangshanensis is the first gallodactylid pterosaur found in Asia. Its discovery not only provides much more osteological information about the Gallodactylidae but also indicates that the ctenochasmatoid pterosaurs were highly diverse in the Early Cretaceous. The filamentous structures preserved near the dorsal and posterior margins of the posterior portion of the skull and around the neck indicate that it had an epidermal covering and may have been a warm-blooded animal.     

Identification of the first postcranial skeleton of Aristonectes Cabrera (Plesiosauroidea,Elasmosauridae) from the upper Maastrichtian of the south-eastern Pacific,based on a bivariate graphic analysis

Cervical vertebrae of elasmosaurids (Sauropterygia, Plesiosauroidea) have proven to be informative even at family level. The present paper uses a previously developed methodology based on bivariate graphic analysis that evaluates the proportion indices of cervical vertebrae in elasmosaurids, which is applied to a well-preserved postcranial specimen of this group recovered from upper Maastrichtian beds of central Chile. The plotting of such indices groups the Chilean material with other specimens from the Late Cretaceous of Argentina and Antarctica, the latter referred to the genus Aristonectes. This result allows us to identify the first relatively complete individual of the genus based on non-cranial characters. The identification of the Chilean specimen as Aristonectes sp. provides important new data on the postcranial skeleton of the genus, that has remained poorly known until now.     

A new species of Tupuxuara (Thalassodromidae,Azhdarchoidea) from the Lower Cretaceous Santana Formation of Brazil,with a note on the nomenclature of Thalassodromidae

A new species of the sail-crested pterosaur Tupuxuara is described from the Santana Formation of Brazil, Tupuxuara deliradamus sp. nov. The holotype, a partial skull, and a larger, partial skull referred to the same taxon differs from Tupuxuara leonardii by having a nasoantorbital fenestra with an acutely-angled posterior border with a long, straight posterodorsal margin, a reclined cranium, and an orbit situated entirely in the ventral half of the nasoantorbitral fenestra. Unfortunately, neither specimen is comparable with the fragmentary rostrum representing Tupuxuara longicristatus. In addition, resolution of a recent nomenclatural problem over the correct name for the clade containing Tupuxuara and its sister taxon, Thalassodromeus, is provided. Both genera are used by different authors as the nomenclatural basis for the group, but “Tupuxuaridae” has never been explicitly erected as a new taxon, and therefore fails to meet ICZN criteria that new taxa are only valid if authors clearly indicate their intention to establish new names. By contrast, “Thalassodrominae” was explicitly erected as a name for the Thalassodromeus + Tupuxuara clade, thereby fulfilling all ICZN requirements for naming of a new taxon and making Thalassodromeus stand as the type genus for this group.     

Reappraisal of Fresnosaurus drescheri (Plesiosauria; Elasmosauridae) from the Maastrichtian Moreno Formation,California, USA

The elasmosaurid Fresnosaurus drescheri, Welles from the contact between the Tierra Loma/Marca members of the Moreno Formation (Maastrichtian), California, USA is reviewed. Most of the features included in Welles's original diagnosis are considered related only to the juvenile ontogenetic stage of the holotype and only specimen. The new diagnosis is based on diagnostic characters of the ilia, including long rectangular shaped sacral facets located in the dorsal part of the shaft, two gentle knobs in the shafts and unexpanded dorsal end. Additional material from the Moreno Formation (numbered under the same number as the F. drescheri holotype but not mentioned by Welles and therefore considered part of a different specimen) are described for the first time. The latter are referred to the aristonectine, being the first evidence of aristonectines from the North Hemisphere.     

A new Triassic pterosaur from Switzerland (Central Austroalpine, Grisons), Raeticodactylus filisurensis gen. et sp. nov.

A new basal non-pterodactyloid pterosaur, Raeticodactylus filisurensis gen. et sp. nov., is reported. It has been discovered in shallow marine sediments from the Upper Triassic of the lowest Kössen beds (late Norian/early Rhaetian boundary) in the central Austroalpine of Canton Grisons (Switzerland). The disarticulated specimen is comprised of an almost complete skull and a partial postcranial skeleton. A high and thin bony, sagittal cranial crest characterizes the anterodorsal region of the skull. The large mandible, with an additional keel-like expansion at the front, partly matches the enlarged sagittal cranial crest. A direct and close relationship to Austriadactylus cristatus, the only known Triassic pterosaur with a bony cranial crest so far, cannot be established. The teeth of the premaxilla are monocuspid and exhibit very strongly bowed enamel wrinkles on the lingual side whereas the enamel is smooth on the labial side. These monocuspid teeth are large and fang-like. The numerous smaller teeth of the maxilla show three, four and five cusps. These are very similar to the teeth of the Triassic pterosaur Eudimorphodon ranzii. The humerus shows a thinner construction than that seen in other Triassic pterosaurs. The femur is quite unusual with a caput femoris perpendicular to the shaft. The bones of the extremities are almost twice as long as the ones from the largest Triassic specimen E. ranzii (MCSNB 2888). The newly described pterosaur is an adult, with a wingspan of approximately 135 cm. A morphofunctional analysis suggests that R. filisurensis was a highly specialized piscivore and possibly a skim-feeder.     

A forefin of Leptonectes solei from the Lower Jurassic (Pliensbachian) of Dorset,UK

An incomplete forefin in the collections of the National Museum of Wales, Cardiff, is herein assigned to the rare leptonectid species, Leptonectes solei, known only from the west Dorset coast. It is only the third specimen of the species and is smaller than both the holotype and referred specimen. The new specimen is from the Lower Jurassic (lower Pliensbachian, Charmouth Mudstone Formation, Tragophylloceras ibex Ammonite Zone, Acanthopleuroceras valdani Ammonite Subzone) of Seatown, Dorset, UK. It confirms that all three species of Leptonectes were present in the Pliensbachian and expands the known diversity of ichthyosaurs for that time period. We show that isolated forefins of Leptonectes can be assigned to a species with a high degree of confidence.     

Kaikaifilu hervei gen. et sp. nov., a new large mosasaur (Squamata,Mosasauridae) from the upper Maastrichtian of Antarctica

We present a large, fragmentary skull and the humerus of a mosasaur (Squamata, Mosasauroidea) recovered from upper Maastrichtian beds of the López de Bertodano Formation in Marambio (=Seymour) Island, Antarctica. The material belongs to a large, adult individual with marked heterodonty as well as unusual humeral features. Different phylogenetic analyses returned the studied specimen within the Tylosaurinae, while the distinctive features of the skull and humerus allow distinguish it from the unique Antarctic known tylosaurine species, Taniwhasaurus antarcticus (Novas et al., 2002), as well as from other known Late Cretaceous mosasaurids from the Southern Hemisphere, thus, justifying the erection of a new taxon, Kaikaifilu hervei gen. et. sp. nov. The different dental types documented in the specimen studied have been previously recorded through isolated teeth from the same locality and were subsequently referred to several genera. This new find is relevant for assessing the previously known fragmentary records of Antarctic mosasaurids, suggesting that its local diversity could be more reduced than previously interpreted. The new material represents the youngest occurrence of tylosaurines in Antarctica.     

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 New Species of Cathayornis from the Lower Cretaceous of Inner Mongolia, China and Its Stratigraphic Significance

An incomplete postcranial avian skeleton is described from the Lower Cretaceous Jingchuan Formation of Otog Banner of western Inner Mongolia and referred to a new species of Cathayornis, C. chabuensis sp. nov. This is the first report of a Cathayornis from outside Liaoning Province. The new discovery indicates that Cathayornis coexisted with Otogornis genghisi, and a more detailed comparison between these two enantiornithine genera shows that Otogornis represents a more primitive genus than Cathayornis. Our analyses further indicate that Cathayornis is an arboreal bird. The discovery of a Cathayornis from this region also confirms that the avian fossil-bearing Jingchuan Formation is comparable to the Jiufotang Formation of the upper Jehol Group in western Liaoning, and should be referred to the middle-late Early Cretaceous.     

A saltasaurine titanosaur (Sauropoda: Titanosauriformes) from the Angostura Colorada Formation (upper Campanian,Cretaceous) of northwestern Patagonia,Argentina

Saltasaurine titanosaurs are characterized by their relatively small size compared to other sauropods, extreme postcranial pneumaticity, and dermal armour covering the body. This group has been reported in the Upper Cretaceous of the Lecho, Allen, and Anacleto formations of Argentina. We describe here a new saltasaurine specimen (MACN-Pv RN 233) from the Campanian of the Angostura Colorada Formation (Río Negro Province) that is represented by eight caudal vertebrae and six osteoderms. This specimen is described in detail and its phylogenetic relationships with the other three known saltasaurines, as well as its implications to the knowledge of caudal vertebra and osteoderm anatomy, are discussed. Our results place MACN-Pv RN 233 more closely related to Saltasaurus loricatus and Rocasaurus muniozi than to Neuquensaurus australis. MACN-Pv RN 233 possesses a combination of features that differ from other saltasaurines, but because of the fragmentary nature of the specimen we decided for the sake of taxonomic stability to not erect a new taxon. This specimen shows the first unambiguous evidence of chevron pneumatisation for a sauropodomorph, implying a broader osteological invasion of the diverticula from the abdominal air sac than previously thought for this group of dinosaurs. MACN-Pv RN 233 preserves two osteoderm morphotypes, one similar to those reported for Neuquensaurus australis and Saltasaurus loricatus. This new specimen expands the distribution of the group to a new geological unit and increases the dinosaur diversity known for the Angostura Colorada Formation.     

A new mawsoniid coelacanth (Actinistia) from the Upper Cretaceous of Southern France

Axelrodichthys megadromos sp. nov. is a coelacanth described based on a single specimen collected in the lower Campanian site of Ventabren motorway, Southern France. The new species is referred to the mawsoniids because of the ornamentation of the skull roof and of the proportionally wide supraorbital series. The specimen belongs to the Mawsonia-Axelrodichthys complex based on features present on the lower jaw and on the basisphenoid. The new species is referred to the genus Axelrodichthys because of proportions of its parietonasal shield and because of the arrangement of the posterior parietals relative to the supraorbitals. Autapomorphic characters, in particular on the parasphenoid, justify the inclusion of the specimen in a new species. The occurrence of a mawsoniid in the Ibero-Amorican Island that formed part of the European Archipelago in the terminal Cretaceous is an evidence of a dispersal event from the southern land masses. The occurrence of A. megadromos in the Campanian-Maastrichtian represents the last occurrence of mawsoniids worldwide, after a gap in the fossil record of about 30 million years. This belated occurrence of Axelrodichthys extends the time range of this genus to approximately 40 myr and suggests that this genus, together with its sister genus Mawsonia, were organisms with a slow morphologic evolution.     

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.