A new ceratopsian dinosaur from the Javelina Formation (Maastrichtian) of West Texas and implications for chasmosaurine phylogeny

Bravoceratops polyphemus gen. et sp. nov. is a large chasmosaurine ceratopsid from the lowermost part of the Javelina Formation (early Maastrichtian) of Big Bend National Park, TX, USA. B. polyphemus has a distinctive narrow snout, a long fenestrate frill, and a fan-shaped median parietal bar with a midline epiparietal on its posterior margin, as well as a symmetrical depression on its dorsal surface at the nexus of the parietal rami. This depression is interpreted to be the attachment point for a second midline epiparietal. This parietal morphology is distinct from that exhibited by Anchiceratops or Pentaceratops. The posterior midline epiparietal in B. polyphemus and its bifurcated quadratojugal–squamosal joint are features shared with the most derived chasmosaurines, Torosaurus and Triceratops. The combination of primitive and derived traits exhibited by B. polyphemus, and its stratigraphic position, is compatible with the gradual transition from basal, to intermediate, to derived chasmosaurines observed throughout the western interior of North America, and with phylogenetic analysis, which suggests that Bravoceratops may be closely related to Coahuilaceratops.     

Turanoceratops tardabilis—the first ceratopsid dinosaur from Asia

Turanoceratops tardabilis from the Upper Cretaceous (Turonian) Bissekty Formation of Dzharakuduk, Uzbekistan, represents the first definite ceratopsid dinosaur recorded from Asia. Reexamination of the original and study of newly collected material indicate that Turanoceratops has well-developed supraorbital horns and apparently lacks a nasal horn. Turanoceratops is more derived than the more or less coeval Zuniceratops from the Moreno Hill Formation of New Mexico in the presence of double-rooted teeth and of two or three teeth in each vertical dental file.     

A new neosuchian with Asian affinities from the Jurassic of northeastern Brazil

Phylogenetic relationships near the origin of extant crocodylians are weakly supported, and this lack of resolution makes for poor estimates of taxonomic and morphological diversity. Previously known taxa are found throughout the Cretaceous in Laurasia and at a few sites from Brazil, Australia, and northern Africa. Here, we report Batrachomimus pastosbonensis gen. et sp. nov. from the Late Jurassic of northeastern Brazil, which is deeply nested within Neosuchia and associated to the Cretaceous Asian paralligatorids. The new taxon predates all other members of Paralligatoridae and its immediate sister group (including Eusuchia) by 30 million years. A nearly complete skull, osteoderms, and limb bones were recovered, and autapomorphies of B. pastosbonensis include a scalloped lateral margin of the rostrum in dorsal view, unsculpted alveolar margin at the caudalmost portion of the maxilla, blunt lateral prongs on the jugal at the base of the postorbital bar, hourglass shaped choanae, and pterygoid choanal septum extended between the palatal shelves of the palatines. The crocodyloid-like skull proportions and the slender, conical teeth suggest an amphibious and piscivorous life history to this 1 m long animal.

     

Ontogenetic data and the evolutionary origin of the mammalian scapula

One of the most useful diagnostic therian features is the division of the lateral surface of the scapula into two fossae separated by a scapular spine. A new model to describe this evolutionary innovation based on ontogenetic data is provided, consistent with information provided by recent fossil discoveries. The development of the scapula in three didelphid and one dasyurid marsupials using histological sections was studied. Only the ventral, acromial portion of the scapular spine, which originates from the anterior margin of the scapular blade, is preformed in cartilage. The major dorsal portion is formed at a later stage by appositional bone, which expands from the perichondral ossification of the scapula into an intermuscular aponeurosis between the supra-and infraspinatus muscles. This intermuscular aponeurosis inserts more or less in the middle of the lateral surface of the developing scapula. Thus, the floor of the supraspinous fossa is already present at the beginning of scapular development, simultaneous with the infraspinous fossa. The portion of the scapular spine that is situated dorsal to the acromial process is hypothesized to be a neomorphic structure of therians. The dorsal portion of the scapular spine evolved as additional attachment for powerful supraspinatus and infraspinatus muscles meeting near the middle of the lateral scapula.     

A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution

Toothless pterosaurs played a key role in broadening the taxonomic, morphological and ecological diversity of Cretaceous pterosaurs. Here we report a complete, articulated skeleton of a 1.4-m-wingspan pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China which is identified as a new genus and species, Shenzhoupterus chaoyangensis gen. et sp. nov. The new taxon is edentulous, with a relatively large skull and a remarkably large, tall nasoantorbital fenestra that extends well above the main part of the braincase. Phylogenetic analysis shows that Shenzhoupterus gen. nov. belongs in a distinct clade of azhdarchoid pterosaurs, formally recognised here as a new family, Chaoyangopteridae, that also includes Chaoyangopterus, Jidapterus and Eoazhdarcho from the Jiufotang Formation and Eopteranodon from the Yixian Formation. These new data clarify recent confusion surrounding the systematics of these Lower Cretaceous taxa and provide new insights into the evolutionary history of pterosaurs.     

A relict trematosauroid (Amphibia: Temnospondyli) from the Middle Jurassic of the Junggar Basin (NW China)

A temnospondyl ilium from the uppermost Toutunhe Formation (Middle Jurassic, Callovian) of the southern Junggar Basin is described. Among the known temnospondyls it is very unusual in morphology because of its very long and slender shaft. It compares closely only to the ilium of one of the latest known trematosaurids from the Ladinian of southern Germany. The Toutunhe Formation has also yielded vertebrae and skull fragments of temnospondyls which belong to the brachyopid Gobiops from the Upper Jurassic of Mongolia. Brachyopoid ilia do not, however, display a morphology similar to that of the new specimen. It is therefore concluded that this specimen represents a second taxon of temnospondyl from the Toutunhe Formation, which probably represents the latest surviving trematosauroid. The Trematosauroidea, which was hitherto exclusively known from the Lower to early Upper Triassic, therefore joins the Brachyopoidea – and possibly the Capitosauroidea – as another group of temnospondyls which survived the end-Triassic mass extinction.     

A Middle Jurassic heterodontosaurid dinosaur from Patagonia and the evolution of heterodontosaurids

Heterodontosauridae is a morphologically divergent group of dinosaurs that has recently been interpreted as one of the most basal clades of Ornithischia. Heterodontosaurid remains were previously known from the Early Jurassic of southern Africa, but recent discoveries and studies have significantly increased the geographical and temporal range for this clade. Here, we report a new ornithischian dinosaur from the Middle Jurassic Ca?adón Asfalto Formation in central Patagonia, Argentina. This new taxon, Manidens condorensis gen. et sp. nov., includes well-preserved craniomandibular and postcranial remains and represents the only diagnostic ornithischian specimen yet discovered in the Jurassic of South America so far. Derived features of its anatomy indicate that Manidens belongs to Heterodontosauridae, as the sister taxon of Heterodontosaurus and other South African heterodontosaurids. The presence of posterior dentary teeth with high crowns but lacking extensive wear facets in Manidens suggests that this form represents an intermediate stage in the development of the remarkable adaptations to herbivory described for Heterodontosaurus. The dentition of Manidens condorensis also has autapomorphies, such as asymmetrically arranged denticles in posterior teeth and a mesially projected denticle in the posteriormost teeth. At an estimated total length of 60?C75?cm, Manidens furthermore confirms the small size of basal heterodontosaurids.     

Associated skeletons of a new middle Triassic "Rauisuchia" from Brazil

For more than 30 million years, in early Mesozoic Pangea, ??rauisuchian?? archosaurs were the apex predators in most terrestrial ecosystems, but their biology and evolutionary history remain poorly understood. We describe a new ??rauisuchian?? based on ten individuals found in a single locality from the Middle Triassic (Ladinian) Santa Maria Formation of southern Brazil. Nine articulated and associated skeletons were discovered, three of which have nearly complete skulls. Along with sedimentological and taphonomic data, this suggests that those highly successful predators exhibited some kind of intraspecific interaction. Other monotaxic assemblages of Triassic archosaurs are Late Triassic (Norian-Rhaetian) in age, approximately 10 million years younger than the material described here. Indeed, the studied assemblage may represent the earliest evidence of gregariousness among archosaurs, adding to our knowledge on the origin of a behavior pattern typical of extant taxa.     

Eocene bunoselenodont Artiodactyla from southern Thailand and the early evolution of Ruminantia in South Asia

Although Asia is thought to have played a critical role in the basal radiation of Ruminantia, the fossil record of early selenodont artiodactyls remains poorly documented in this region. Dental remains of a new bunoselenodont artiodactyl are described from the late Eocene of Krabi, southern Thailand. This new form, Krabitherium waileki gen. et sp. nov, is tentatively referred to the Tragulidae (Ruminantia) on the basis of several dental features, including a weak Tragulus fold and the presence of a deep groove on the anterior face of the entoconid. Although this new form is suggestive of the enigmatic ?Gelocus gajensis Pilgrim 1912 from the “base of the Gaj” (lower Chitarwata Formation) of the Bugti Hills (Central Pakistan), K. waileki most likely represents an early representative of a relatively bunodont group of tragulids that includes the genus Dorcabune, known from the Miocene of south Asia. This addition to the Eocene record of early ruminants attests to the antiquity of the group in Southeast Asia and lends support to the hypothesis that the Tragulidae represents one of the first offshoots in the evolutionary history of Ruminantia.     

A new basal galeomorph shark (Synechodontiformes, Neoselachii) from the Early Jurassic of Europe

Palaeospinacids are a group of basal galeomorph sharks and are placed in the order Synechodontiformes (Chondrichthyes, Neoselachii) ranging from the Permian to the Eocene. Currently, there is a controversy concerning the identity of diagnostic characters for distinguishing palaeospinacid genera because of very similar dental morphologies and the scarcity of articulated skeletal material. The most notable character for distinguishing species within the Palaeospinacidae is, however, the dental morphology. The main dental character uniting all palaeospinacids is the very specialised pseudopolyaulacorhize root vascularisation. A re-examination of articulated neoselachian skeletons from the Lower Jurassic of Lyme Regis (England) and Holzmaden (S Germany), and recently discovered specimens from the Upper Jurassic of the Solnhofen area and Nusplingen (S Germany) has yielded several hitherto unrecognised complete skeletons of the palaeospinacids Synechodus and Paraorthacodus enabling a re-evaluation of characters. These specimens indicate that the number of dorsal fins and the presence or absence of dorsal fin spines represent important features for identifying palaeospinacids. Synechodus bears two dorsal fins without fin spines, whereas Paraorthacodus only has a single dorsal fin lacking a fin spine directly in front of the caudal fin. All palaeospinacids from the Early Jurassic have two spines supporting the dorsal fins and are consequently assigned to a new genus, Palidiplospinax nov. gen. Three species are placed into the new taxon: Synechodus enniskilleni, S. occultidens and S. smithwoodwardi.     

Brainy stuff of long-gone dogs: a reappraisal of the supposed Canis endocranial cast from the Pliocene of Poland

The pre-Quaternary fossil record of Canis in the Old World is scarce, and the first appearance of this genus in Europe remains an enigma. Amongst the oldest fossils assigned to this genus, there is a natural cast of the brain (endocast) collected in W??e 1, Poland, from Pliocene deposits dated between 3.3 and 4.0 Ma. We reexamined this specimen and found that it differs from the brain of Canis in having its region medial to the coronal sulcus heart-shaped in dorsal view, its region rostral to the presylvian sulcus shorter and less constricted laterally, and its cerebellum less overlapped by the cerebrum and lacking a lateral twist of the posterior vermis. We identified this fossil, as well as another fossil canid endocast from W??e 1, as representing the raccoon dog genus Nyctereutes. The previously reported presence of Canis in W??e 1 is therefore not confirmed. Specifically, both endocasts can be referred to N. donnezani because this is the only species of Nyctereutes that has been recognised in this locality on the basis of craniomandibular and dental fossils. Our study represents a taxonomic application of comparative neuroanatomical and palaeoneurological data, an approach that may become increasingly useful with the growing knowledge of the endocranial morphology of fossil mammals.     

A new scincomorph lizard from the Palaeocene of Belgium and the origin of Scincoidea in Europe

Well-preserved jaw fragments and skull bones of a new scincomorphan lizard are described from the continental Hainin Formation (Middle Palaeocene) of the Mons Basin in southwestern Belgium. This taxon, Scincoideus haininensis gen. et sp. nov., is characterised by vermiculate dermal rugosities and represents the earliest formally recognised scincoid in Europe. It differs from cordylids (girdled lizards) by lacking the parietal downgrowth and differs from scincids (skinks) by having robustly built bones and a well-developed lingual cusp on tooth crowns. By its mosaic characters, S. haininensis is considered here as being a basal scincoid lizard close to the origin of cordylids and scincids. Palaeobiogeographic analysis suggests that scincoids were present in Europe from the Palaeocene or maybe even before the Cretaceous/Palaeogene boundary and that they were replaced by modern scincoids such as true scincids and cordylids during the Eocene. The origin of the latter two families in Europe was very likely related with the immigration of the first modern mammals.Electronic supplementary material  Supplementary material is available for this article at and accessible for authorised users.     

Artiodactyls from the Pondaung Formation (Myanmar): new data and reevaluation of the South Asian Faunal Province during the Middle Eocene

Although Asia is thought to have played a critical role in the radiation of artiodactyls, the fossil record of stem selenodonts (“dichobunoids”) remains dramatically poor in tropical Asian regions. In this study, we report a new dichobunid genus and species Cadutherium kyaukmagyii and a new basal ruminant genus and species Irrawadymeryx pondaungi, from the late Middle Eocene Pondaung Formation, Central Myanmar. Although the scarcity of the present material prevents any attempts to investigate the phylogenetic relationships of Cadutherium with contemporaneous forms from other Holarctic landmasses, this new form shed new light on the diversity of these small rabbit-like ungulates during a key period of their evolutionary history. Reexamination of the small-bodied artiodactyls from Pondaung leads us to propose new identifications of certain published specimens and, in turn, to investigate the temporal and geographic distribution of taxa recognized in the Pondaung Formation. Although fragmentary, these potential new taxa reveal an unsuspected diversity of small forms among artiodactyls of Pondaung. This addition to the Eocene record of dichobunoids and early ruminants provides further insight in the diversity of dental patterns among small artiodactyls from the Pondaung Formation and attests to the antiquity of these groups in Southeast Asia.     

A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America

Dromaeosaurids from the Maastrichtian of North America have a poor fossil record and are known largely from isolated teeth, which have typically been referred to taxa based on more complete material from earlier Campanian strata. An almost complete maxilla with well-preserved dentition and an associated dentary from the Hell Creek Formation of Montana are used to establish a new dromaeosaurid taxon in the latest Maastrichtian, immediately prior to the end-Cretaceous extinction event. Acheroraptor temertyorum gen. et sp. nov. is differentiated from other dromaeosaurids on the basis of a hypertrophied postantral wall that projects posteriorly into the antorbital fenestra, a maxillary fenestra positioned low in the antorbital fossa and directly posterior to the promaxillary fenestra, and distinctive dentition with marked apicobasal ridges. The new material allows a dromaeosaurid from the Maastrichtian of North America to be placed within a phylogenetic framework for the first time. Phylogenetic analysis suggests Acheroraptor is a velociraptorine that is more closely related to Asian dromaeosaurids, including Tsaagan and Velociraptor, than it is to Dromaeosaurus, Saurornitholestes, or any other taxon from North America. As part of the Lancian Tyrannosaurus–Triceratops fauna, A. temertyorum is the latest occurring dromaeosaurid. Its relationships and occurrence suggest a complex historical biogeographic scenario that involved multiple, bi-directional faunal interchanges between Asia and North America during the Late Cretaceous.     

Osteological evidence for sister group relationship between pseudo-toothed birds (Aves: Odontopterygiformes) and waterfowls (Anseriformes)

The phylogenetic affinities of the extinct pseudo-toothed birds have remained controversial. Some authors noted that they resemble both pelicans and allies (Pelecaniformes) and tube-nosed birds (Procellariiformes), but assigned them to a distinct taxon, the Odontopterygiformes. In most recent studies, the pseudo-toothed birds are referred to the family Pelagornithidae inside the Pelecaniformes. Here, I perform a cladistic analysis with five taxa of the pseudo-toothed birds including two undescribed new species from the Early Tertiary of Morocco. The present hypothesis strongly supports a sister group relationship of pseudo-toothed birds (Odontopterygiformes) and waterfowls (Anseriformes). The Odontoanserae (Odontopterygiformes plus Anseriformes) are the sister group of Neoaves. The placement of the landfowls (Galliformes) as the sister taxon of all other neognathous birds does not support the consensus view that the Galloanserae (Galliformes plus Anseriformes) are monophyletic. Electronic supplementary material  Supplementary material is available in the online version of this article at . Communicated by G. Mayr     

An unusual archosaurian from the marine Triassic of China

A new Triassic archosaurian from China shows a number of aquatic specializations, of which the most striking is the extreme lateral compression of the long tail. Others that may also reflect aquatic adaptations include platelike scapula and coracoid, elongate neck with extremely long and slender ribs, and reduction of osteoderms. In contrast, its pelvic girdle and hind limb have no aquatic modifications. Anatomic features, taphonomy, and local geological data suggest that it may have lived in a coastal–island environment. This lifestyle, convergent with some Jurassic marine crocodyliforms that lived at least 40 million years later and the saltwater species of extant Crocodylus, contradicts with the prevailing view that Triassic archosaurians were restricted to nonmarine ecosystems. Its mosaic anatomy represents a previously unknown ecomorph within primitive archosaurians.Electronic Supplementary Material  Supplementary material is available for this article at     

Gaudeamus lavocati sp. nov. (Rodentia, Hystricognathi) from the early Oligocene of Zallah, Libya: first African caviomorph?

A new African species of hystricognathous rodent, Gaudeamus lavocati sp. nov., is described herein from the early Oligocene deposits of Zallah locality (Sirt basin, Central Libya). The dental morphology of this species is very close to that of some earliest South American caviomorphs. It allows a reinterpretation of molar crest homologies among earliest caviomorphs, pentalophodonty being confirmed as the plesiomorphic molar condition in Caviomorpha. This morphological resemblance argues for close affinities between Gaudeamus and earliest South American hystricognaths. Cladistic analysis supports Gaudeamus lavocati sp. nov. as the first known African representative of Caviomorpha, implying that its ancestors were part of the African phiomyid group that crossed the South Atlantic by a direct immigration route. Alternatively, the series of derived dental features of Gaudeamus could also be interpreted as evolutionary synchronous convergences of an African hystricognath lineage towards the specialized pattern of some caviomorphs. However, the high level of similarities concerning teeth morphology and enamel microstructure and the similar age of fossiliferous strata on both continents make this interpretation less probable. The phylogenetic position of this taxon is of considerable importance because it represents an enigmatic component of the phiomorph–caviomorph radiation in Africa and appears as a new clue toward the understanding of caviomorph origins.     

Evidence for high taxonomic and morphologic tyrannosauroid diversity in the Late Cretaceous (Late Campanian) of the American Southwest and a new short-skulled tyrannosaurid from the Kaiparowits formation of Utah

The fossil record of late Campanian tyrannosauroids of western North America has a geographic gap between the Northern Rocky Mountain Region (Montana, Alberta) and the Southwest (New Mexico, Utah). Until recently, diagnostic tyrannosauroids from the Southwest were unknown until the discovery of Bistahieversor sealeyi from the late Campanian of New Mexico. Here we describe an incomplete skull and postcranial skeleton of an unusual tyrannosaurid from the Kaiparowits Formation (Late Cretaceous) of Utah that represents a new genus and species, Teratophoneus curriei. Teratophoneus differs from other tyrannosauroids in having a short skull, as indicated by a short and steep maxilla, abrupt angle in the postorbital process of the jugal, laterally oriented paroccipital processes, short basicranium, and reduced number of teeth. Teratophoneus is the sister taxon of the Daspletosaurus + Tyrannosaurus clade and it is the most basal North American tyrannosaurine. The presence of Teratophoneus suggests that dinosaur faunas were regionally endemic in the west during the upper Campanian. The divergence in skull form seen in tyrannosaurines indicates that the skull in this clade had a wide range of adaptive morphotypes.     

A juvenile lizard specimen with well-preserved skin impressions from the Upper Jurassic/Lower Cretaceous of Daohugou, Inner Mongolia, China

Lizards are now relatively well known from the Yixian Formation of northeastern China. In this study, we describe a juvenile lizard from a fossil horizon at Daohugou, Inner Mongolia. These beds predate the Yixian Formation, and are probably Late Jurassic or earliest Cretaceous in age. The new specimen thus documents the first lizard material from the Daohugou locality and is the earliest lizard skeleton from China. Comparisons with developmental stages of modern lizards suggest the Daohugou lizard is a hatchling. Although tiny, the specimen is notable in preserving exquisite skin impressions showing the variation in scalation across the body, the shape and position of the cloacal outlet, and details of the manus and pes. These are the earliest recorded lepidosaurian skin traces. In its general proportions and the possession of paired frontals, the small Daohugou lizard resembles both the Yixian taxon Yabeinosaurus tenuis and the questionable Jeholacerta formosa, but it differs from the latter in scalation and, based on other characters, may be distinct from both.