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A fossil‐bearing locality near Padhri village, Dhok Pathan, 55 km away from the tehsil Dina, Jhelum District, in the Potwar Plateau, Middle Siwaliks, Punjab, northern Pakistan, is significantly rich in mammalian fossils. This site has provided an abundant mammalian fossil fauna of Late Miocene age from the Dhok Pathan Formation (Fm.). The recovered material belongs to four families: Equidae (horses), Rhinocerotidae (rhinos), Bovidae (cows), and Suidae (pigs). We discovered a new skull of hipparionine Hipparion theobaldi from this locality along with 22 specimens from the associated assemblage of fossil mammals. The recovered material includes seven other species: the aceratheriine Chilotherium intermedium, boselaphines Tragoportax punjabicus, Selenoportax vexillarius, Pachyportax latidens, the antelope Gazella lydekkeri and suinine Propotamochoerus hysudricus. The specimens are isolated teeth, fragments of maxilla, mandibles and horn cores. The Dhok Pathan Fm. is generally composed of claystone, siltstone and sandstone beds and, based on the mammalian fauna, the Padhri fossil locality is dated as Late Miocene. Thi99s formation was deposited in a subtropical paleoenvironment and the predominance of fossil bovids indicates extremely moist conditions with small but frequent standing water bodies. [ABSTRACT FROM AUTHOR]

Saturnalia tupiniquim is a sauropodomorph dinosaur from the Late Triassic (Carnian–c. 233 Ma) Santa Maria Formation of Brazil. Due to its phylogenetic position and age, it is important for studies focusing on the early evolution of both dinosaurs and sauropodomorphs. The osteology of Saturnalia has been described in a series of papers, but its cranial anatomy remains mostly unknown. Here, we describe the skull bones of one of its paratypes (only in the type-series to possess such remains) based on CT Scan data. The newly described elements allowed estimating the cranial length of Saturnalia and provide additional support for the presence of a reduced skull (i.e. two thirds of the femoral length) in this taxon, as typical of later sauropodomorphs. Skull reduction in Saturnalia could be related to an increased efficiency for predatory feeding behaviour, allowing fast movements of the head in order to secure small and elusive prey, a hypothesis also supported by data from its tooth and brain morphology. A principal co-ordinates analysis of the sauropodomorph jaw feeding apparatus shows marked shifts in morphospace occupation in different stages of the first 30 million years of their evolutionary history. One of these shifts is observed between non-plateosaurian and plateosaurian sauropodomorphs, suggesting that, despite also having an omnivorous diet, the feeding behaviour of some early Carnian sauropodomorphs, such as Saturnalia, was markedly different from that of later Triassic taxa. A second shift, between Late Triassic and Early Jurassic taxa, is congruent with a floral turnover hypothesis across the Triassic-Jurassic boundary. [ABSTRACT FROM AUTHOR]

We here describe the first fossil remains of a green lizard of the Lacerta group from the late Miocene (MN 13) of the Solnechnodolsk locality in southern European Russia. This region of Europe is crucial for our understanding of the paleobiogeography and evolution of these middle-sized lizards. Although this clade has a broad geographical distribution across the continent today, its presence in the fossil record has only rarely been reported. In contrast to that, the material described here is abundant, consists of a premaxilla, maxillae, frontals, parietals, jugals, quadrate, pterygoids, dentaries and vertebrae. The comparison of these elements to all extant green lizard species shows that these fossils are indistinguishable from Lacerta trilineata. Thus, they form the first potential evidence of the occurrence of this species in the Miocene. This may be also used as a potential calibration point for further studies. Together with other lizard fossils, Solnechnodolsk shows an interesting combination of survivors and the dawn of modern species. This locality provides important evidence for the transition of an archaic Miocene world to the modern diversity of lizards in Europe. In addition, this article represents a contribution to the knowledge of the comparative osteological anatomy of the selected cranial elements in lacertids. This study gives special emphasis to the green lizards, but new data are also presented for related taxa, e.g., Timon lepidus, Podarcis muralis or Zootoca vivipara. Although the green lizards include several cryptic species for which determination based on isolated osteological material would be expected to be difficult, our comparisons show several important morphological differences, although a high degree of variability is present. [ABSTRACT FROM AUTHOR]

We describe a new probainognathian cynodont, Pseudotherium argentinus, from the early Late Triassic Ischigualasto Formation of Argentina. Pseudotherium adds to a growing assemblage of small Triassic cynodonts that offers new insight into events leading up to the origin of crown Mammalia and the successively more inclusive Mammaliaformes and Mammaliamorpha. Using high-resolution X-ray computed tomography, we illustrate and describe the holotype and only known specimen, which consists of a well-preserved isolated skull. It preserves apomorphic features of the orbit and braincase. Prefrontal and vestigial postorbital bones are present, despite the absence of an ossified postorbital bar. As in Brasilitherium riograndensis, thin turbinal-like bones are present in the nasopharyngeal passage, and we discuss impediments to establishing their identity and function. Compared to more basal cynodonts, the cochlea is elongated but uncoiled and in this and other features it resembles basal mammaliamorphs. Our analysis found weak support for Pseudotherium as the sister taxon of Tritylodontidae. However, a broader assessment of its relationships in light of additional character data from the literature and unpublished computed tomography data suggest that it may be more realistic to view the relationships of Pseudotherium as an unresolved polytomy with tritylodontids, and the taxa referred to as tritheledontids and brasilodontids (groups of variable membership and questionable monophyly). Thus, Pseudotherium may lie just inside or just outside of Mammaliamorpha, and there is also weak character support for its sister taxon relationship with Brasilitherium. Our results amplify previous conclusions that phylogenetic relationships in this adaptive radiation of small cynodonts will remain somewhat uncertain until more complete specimens are recovered, and until high-resolution CT scans of existing specimens become available to the larger community. Toward that goal, we make the CT dataset for the holotype of Pseudotherium argentinus publically available under a Creative Commons license at . [ABSTRACT FROM AUTHOR]

A new genus and species of non-hadrosaurid hadrosauroid, Gobihadros mongoliensis, is described from a virtually complete and undeformed skull and postcranial skeleton, as well as extensive referred material, collected from the Baynshire Formation (Cenomanian-Santonian) of the central and eastern Gobi Desert, Mongolia. Gobihadros mongoliensis is the first non-hadrosaurid hadrosauroid from the Late Cretaceous of central Asia known from a complete, articulated skull and skeleton. The material reveals the skeletal anatomy of a proximate sister taxon to Hadrosauridae in remarkable detail. Gobihadros is similar to Bactrosaurus johnsoni and Gilmoreosaurus mongoliensis, but can be distinguished from them in several autapomorphic traits, including the maximum number (three) of functional dentary teeth per tooth position, a premaxillary oral margin with a ‘double-layer morphology’, and a sigmoidal dorsal outline of the ilium with a well-developed, fan-shaped posterior process. All of these characters in Gobihadros are inferred to be convergent in Hadrosauridae. Phylogenetic analysis positions Gobihadros mongoliensis as a Bactrosaurus-grade hadrosauromorph hadrosauroid. Its relationship with Maastrichtian hadrosaurids from Asia (e.g., Saurolophus angustirostris, Kerberosaurus manakini, Wulagasaurus dongi, Kundurosaurus nagornyi) are sufficiently distant to indicate that these latter taxa owe their distribution to migration from North America across Beringia, rather than having a common Asian origin with Go. mongoliensis. [ABSTRACT FROM AUTHOR]

A braincase of the Cretaceous titanosaurian sauropod Malawisaurus dixeyi, complete except for the olfactory region, was CT scanned and a 3D rendering of the endocast and inner ear was generated. Cranial nerves appear in the same configuration as in other sauropods, including derived features that appear to characterize titanosaurians, specifically, an abducens nerve canal that passes lateral to the pituitary fossa rather than entering it. Furthermore, the hypoglossal nerve exits the skull via a single foramen, consistent with most titanosaurians, while other saurischians, including the basal titanosauriform, Giraffatitan, contain multiple rootlets. The size of the vestibular labyrinth is smaller than in Giraffatitan, but larger than in most derived titanosaurians. Similar to the condition found in Giraffatitan, the anterior semicircular canal is larger than the posterior semicircular canal. This contrasts with more derived titanosaurians that contain similarly sized anterior and posterior semicircular canals, congruent with the interpretation of Malawisaurus as a basal titanosaurian. Measurements of the humerus of Malawisaurus provide a body mass estimate of 4.7 metric tons. Comparison of body mass to radius of the semicircular canals of the vestibular labyrinth reveals that Malawisaurus fits the allometric relationship found in previous studies of extant mammals and Giraffatitan brancai. As in Giraffatitan, the anterior semicircular canal is significantly larger than is predicted by the allometric relationship suggesting greater sensitivity and slower movement of the head in the sagittal plane. [ABSTRACT FROM AUTHOR]

Convoluted nasal passages are an enigmatic hallmark of Ankylosauria. Previous research suggested that these convoluted nasal passages functioned as heat exchangers analogous to the respiratory turbinates of mammals and birds. We tested this hypothesis by performing a computational fluid dynamic analysis on the nasal passages of two ankylosaurs: Panoplosaurus mirus and Euoplocephalus tutus. Our models predicted that Panoplosaurus and Euoplocephalus would have required 833 and 1568 thermal calories, respectively, to warm a single breath of air by 20°C. Heat recovery during exhalation resulted in energy savings of 65% for Panoplosaurus and 84% for Euoplocephalus. Our results fell well within the range of values for heat and water savings observed in extant terrestrial amniotes. We further tested alternate airway reconstructions that removed nasal passage convolutions or reduced nasal vestibule length. Our results revealed that the extensive elaboration observed in the nasal vestibules of ankylosaurs was a viable alternative to respiratory turbinates with regards to air conditioning. Of the two dinosaurs tested, Euoplocephalus repeatedly exhibited a more efficient nasal passage than Panoplosaurus. We suggest that the higher heat loads associated with the larger body mass of Euoplocephalus necessitated these more efficient nasal passages. Our findings further indicate that the evolution of complicated airways in dinosaurs may have been driven by the thermal requirements of maintaining cerebral thermal homeostasis. [ABSTRACT FROM AUTHOR]

The great diversity of dinosaurian tooth shapes and sizes, and in particular, the amazing dental complexity in derived ornithischians has attracted a lot of attention. However, the evolution of dental batteries in hadrosaurids and ceratopsids is difficult to understand without a broader comparative framework. Here we describe tooth histology and development in the "middle" Cretaceous ornithischian dinosaur Changchunsaurus parvus, a small herbivore that has been characterized as an early ornithopod, or even as a more basal ornithischian. We use this taxon to show how a "typical" ornithischian dentition develops, copes with wear, and undergoes tooth replacement. Although in most respects the histological properties of their teeth are similar to those of other dinosaurs, we show that, as in other more derived ornithischians, in C. parvus the pulp chamber is not invaded fully by the newly developing replacement tooth until eruption is nearly complete. This allowed C. parvus to maintain an uninterrupted shearing surface along a single tooth row, while undergoing continuous tooth replacement. Our histological sections also show that the replacement foramina on the lingual surfaces of the jaws are likely the entry points for an externally placed dental lamina, a feature found in many other ornithischian dinosaurs. Surprisingly, our histological analysis also revealed the presence of wavy enamel, the phylogenetically earliest occurrence of this type of tissue. This contradicts previous interpretations that this peculiar type of enamel arose in association with more complex hadrosauroid dentitions. In view of its early appearance, we suggest that wavy enamel may have evolved in association with a shearing-type dentition in a roughly symmetrically-enameled crown, although its precise function still remains somewhat of a mystery. [ABSTRACT FROM AUTHOR]

Paleopathologies document skeletal damage in extinct organisms and can be used to infer the causes of injury, as well as aspects of related biology, ecology and behavior. To date, few studies have been undertaken on Jurassic marine reptiles, while ichthyosaur pathologies in particular have never been systematically evaluated. Here we survey 41 specimens of the apex predator ichthyosaur Temnodontosaurus from the Early Jurassic of southern Germany in order to document the range and absolute frequency of pathologies observed in this taxon as a function of the number of specimens examined. According to our analysis, most observed pathologies in Temnodontosaurus are force-induced traumas with signs of healing, possibly inflicted during aggressive interactions with conspecifics. When the material is preserved, broken ribs are correlated in most of the cases with traumas elsewhere in the skeleton such as cranial injuries. The range of cranial pathologies in Temnodontosaurus is similar to those reported for extinct cetaceans and mosasaurs, which were interpreted as traces of aggressive encounters. Nevertheless, Temnodontosaurus differs from these other marine amniotes in the absence of pathologies in the vertebral column, consistent with the pattern previously documented in ichthyosaurs. We did not detect any instances of avascular necrosis in Temnodontosaurus from southern Germany, which may reflect a shallow diving life style. This study is intended to provide baseline data for the various types of observed pathologies in large ichthyosaurs occupying the ‘apex predator’ niche, and potentially clarifies aspects of species-specific behavior relative to other ichthyosaurs and marine amniotes. [ABSTRACT FROM AUTHOR]

The soft-tissue reconstruction and associated osteology of the North American therizinosaurian Nothronychus mckinleyi is updated. The cranial nerve topology is revised, bringing it more in line with coelurosaurs. The trunk of the trigeminal nerve is very short, with an incompletely intracranial trigeminal ganglion, an ophthalmic branch diverging anteriorly first, with later divergences of the maxillomandibular branches, following typical pathways. The facial nerve has been re-evaluated, resulting in a very typical configuration with an extracranial geniculate ganglion. The single foramen leading to the cochlea probably transmitted the vestibulocochlear nerve, along with some fibers of the facial. This configuration is reduced from the more standard three foramina (vestibular, cochlear, and facial) and may be apomorphic for therizinosaurs. Some alteration is proposed for the dorsiflexive musculature. The insertion point for m. transversospinalis capitis is partially changed to extend onto the parietal, along with a proposed functional difference in the moment arm. The expansion of the basicranial pneumatic system is limited to the paratympanic system, enhancing low frequency sound sensitivity. There is little expansion of the median pharyngeal and subcondylar sinuses. Ossification of the surrounding epithelium may provide some information on the embryology of the theropod skull. It may be associated with a reduced stress field, or the general similarity of the basicranium with anterior cervical vertebrae may reflect activation of a cervical vertebral (Hox) gene regulating ossification of the pneumatic sinuses. This might be a local, selectively neutral, fixed gene in the basicranium reflecting embryological regulation of cervical vertebrae development. [ABSTRACT FROM AUTHOR]

The largest reported ichthyosaurs lived during the Late Triassic (~235–200 million years ago), and isolated, fragmentary bones could be easily mistaken for those of dinosaurs because of their size. We report the discovery of an isolated bone from the lower jaw of a giant ichthyosaur from the latest Triassic of Lilstock, Somerset, UK. It documents that giant ichthyosaurs persisted well into the Rhaetian Stage, and close to the time of the Late Triassic extinction event. This specimen has prompted the reinterpretation of several large, roughly cylindrical bones from the latest Triassic (Rhaetian Stage) Westbury Mudstone Formation from Aust Cliff, Gloucestershire, UK. We argue here that the Aust bones, previously identified as those of dinosaurs or large terrestrial archosaurs, are jaw fragments from giant ichthyosaurs. The Lilstock and Aust specimens might represent the largest ichthyosaurs currently known. [ABSTRACT FROM AUTHOR]

Nothosaurs form a subclade of the secondarily marine Sauropterygia that was well represented in late Early to early Late Triassic marine ecosystems. Here we present and discuss the internal skull anatomy of the small piscivorous nothosaur Nothosaurus marchicus from coastal to shallow marine Lower Muschelkalk deposits (Anisian) of Winterswijk, The Netherlands, which represents the oldest sauropterygian endocast visualized to date. The cranial endocast is only partially encapsulated by ossified braincase elements. Cranial flattening and lateral constriction by hypertrophied temporal musculature grant the brain a straight, tubular geometry that lacks particularly well-developed cerebral lobes but does potentially involve distinguishable optic lobes, suggesting vision may have represented an important sense during life. Despite large orbit size, the circuitous muscular pathway linking the basisphenoidal and orbital regions indicates poor oculomotor performance. This suggests a rather fixed ocular orientation, although eye placement and neck manoeuvrability could have enabled binocular if not stereoscopic vision. The proportionally large dorsal projection of the braincase endocast towards the well-developed pineal foramen advocates substantial dependence on the corresponding pineal system in vivo. Structures corroborating keen olfactory or acoustic senses were not identified. The likely atrophied vomeronasal organ argues against the presence of a forked tongue in Nothosaurus, and the relative positioning of external and internal nares contrasts respiratory configurations proposed for pistosauroid sauropterygians. The antorbital domain furthermore accommodates a putative rostral sensory plexus and pronounced lateral nasal glands that were likely exapted as salt glands. Previously proposed nothosaurian ‘foramina eustachii’ arose from architectural constraints on braincase development rather than representing functional foramina. Several modifications to brain shape and accessory organs were achieved through heterochronic development of the cranium, particularly the braincase. In summary, the cranium of Nothosaurus marchicus reflects important physiological and neurosensory adaptations that enabled the group’s explosive invasion of shallow marine habitats in the late Early Triassic. [ABSTRACT FROM AUTHOR]

Fossil sites from Brazil have yielded specimens of spinosaurid theropods, among which the most informative include the cranial remains of Irritator, Angaturama, and Oxalaia. In this work some of their craniodental features are reinterpreted, providing new data for taxonomic and evolutionary issues concerning this particular clade of dinosaurs. The mesial-most tooth of the left maxilla of the holotype of Irritator is regarded as representing the third tooth position, which is also preserved in the holotype of Angaturama. Thus, both specimens cannot belong to the same individual, contrary to previous assumptions, although they could have been the same taxon. In addition, the position of the external nares of Irritator is more comparable to those of Baryonyx and Suchomimus instead of other spinosaurine spinosaurids. In fact, with regards to some craniodental features, Brazilian taxa represent intermediate conditions between Baryonychinae and Spinosaurinae. Such a scenario is corroborated by our cladistic results, which also leave open the possibility of the former subfamily being non-monophyletic. Furthermore, the differences between spinosaurids regarding the position and size of the external nares might be related to distinct feeding habits and degrees of reliance on olfaction. Other issues concerning the evolution and taxonomy of Spinosauridae require descriptions of additional material for their clarification. [ABSTRACT FROM AUTHOR]

The cranial anatomy of the flat-skulled hadrosaurine Edmontosaurus regalis (Ornithischia: Hadrosauridae) is extensively described here, based on the holotype and paratype collected from the middle part of the Horseshoe Canyon Formation in southern Alberta. Focus is given to previously undocumented features of ontogenetic and phylogenetic importance. This description facilitates overall osteological comparisons between E. regalis and other hadrosaurids (especially E. annectens), and revises the diagnosis of E. regalis, to which a new autapomorphy (the dorsal half of the jugal anterior process bearing a sharp posterolateral projection into the orbit) is added. We consider the recently named Ugrunaaluk kuukpikensis from the upper Campanian/lower Maastrichtian of Alaska a nomen dubium, and conservatively regard the Alaskan material as belonging to Edmontosaurus sp.. A phylogenetic analysis of Hadrosauroidea using maximum parsimony further corroborates the sister-taxon relationship between E. regalis and E. annectens. In the strict consensus tree, Hadrosaurus foulkii occurs firmly within the clade comprising all non-lambeosaurine hadrosaurids, supporting the taxonomic scheme that divides Hadrosauridae into Hadrosaurinae and Lambeosaurinae. Within Edmontosaurini, Kerberosaurus is posited as the sister taxon to the clade of Shantungosaurus + Edmontosaurus. The biogeographic reconstruction of Hadrosaurinae in light of the time-calibrated cladogram and probability calculation of ancestral areas for all internal nodes reveals a significantly high probability for the North American origin of the clade. However, the Laramidia–Appalachia dispersals around the Santonian–Campanian boundary, inferred from the biogeographic scenario for the North American origin of Hadrosaurinae, are in conflict with currently accepted paleogeographic models. By contrast, the Asian origin of Hadrosaurinae with its relatively low probability resulting from the biogeographic analysis is worth seriously considering, despite the lack of fossil material from the Santonian and lower Campanian of Asia. Extra fossil collecting in appropriate geographic locations and stratigraphic intervals of Asia and Europe will help to clarify the biogeographic dynamics of hadrosaurine dinosaurs in the near future. [ABSTRACT FROM AUTHOR]

Euchambersia mirabilis is an iconic species of Permo-Triassic therapsid because of its unusually large external maxillary fossa linked through a sulcus to a ridged canine. This anatomy led to the commonly accepted conclusion that the large fossa accommodated a venom gland. However, this hypothesis remains untested so far. Here, we conducted a μCT scan assisted reappraisal of the envenoming capacity of Euchambersia, with a special focus on the anatomy of the maxillary fossa and canines. This study shows that the fossa, presumably for the venom-producing gland, is directly linked to the maxillary canal, which carries the trigeminal nerve (responsible for the sensitivity of the face). The peculiar anatomy of the maxillary canal suggests important reorganisation in the somatosensory system and that a ganglion could possibly have been present in the maxillary fossa instead of a venom gland. Nevertheless, the venom gland hypothesis is still preferred since we describe, for the first time, the complete crown morphology of the incisiform teeth of Euchambersia, which strongly suggests that the complete dentition was ridged. Therefore Euchambersia manifests evidence of all characteristics of venomous animals: a venom gland (in the maxillary fossa), a mechanism to deliver the venom (the maxillary canal and/or the sulcus located ventrally to the fossa); and an apparatus with which to inflict a wound for venom delivery (the ridged dentition). [ABSTRACT FROM AUTHOR]

We describe two new cynodonts from the early Late Triassic of southern Brazil. One taxon, Bonacynodon schultzi gen. et sp. nov., comes from the lower Carnian Dinodontosaurus AZ, being correlated with the faunal association at the upper half of the lower member of the Chañares Formation (Ischigualasto-Villa Unión Basin, Argentina). Phylogenetically, Bonacynodon is a closer relative to Probainognathus jenseni than to any other probainognathian, bearing conspicuous canines with a denticulate distal margin. The other new taxon is Santacruzgnathus abdalai gen. et sp. nov. from the Carnian Santacruzodon AZ. Although based exclusively on a partial lower jaw, it represents a probainognathian close to Prozostrodon from the Hyperodapedon AZ and to Brasilodon, Brasilitherium and Botucaraitherium from the Riograndia AZ. The two new cynodonts and the phylogenetic hypothesis presented herein indicate the degree to which our knowledge on probainognathian cynodonts is incomplete and also the relevance of the South American fossil record for understanding their evolutionary significance. The taxonomic diversity and abundance of probainognathians from Brazil and Argentina will form the basis of deep and complex studies to address the evolutionary transformations of cynodonts leading to mammals. [ABSTRACT FROM AUTHOR]

Choerosaurus dejageri, a non-mammalian eutheriodont therapsid from the South African late Permian (~259 Ma), has conspicuous hemispheric cranial bosses on the maxilla and the mandible. These bosses, the earliest of this nature in a eutheriodont, potentially make C. dejageri a key species for understanding the evolutionary origins of sexually selective behaviours (intraspecific competition, ritualized sexual and intimidation displays) associated with cranial outgrowths at the root of the clade that eventually led to extant mammals. Comparison with the tapinocephalid dinocephalian Moschops capensis, a therapsid in which head butting is strongly supported, shows that the delicate structure of the cranial bosses and the gracile structure of the skull of Choerosaurus would be more suitable for display and low energy combat than vigorous head butting. Thus, despite the fact that Choerosaurus is represented by only one skull (which makes it impossible to address the question of sexual dimorphism), its cranial bosses are better interpreted as structures involved in intraspecific selection, i.e. low-energy fighting or display. Display structures, such as enlarged canines and cranial bosses, are widespread among basal therapsid clades and are also present in the putative basal therapsid Tetraceratops insignis. This suggests that sexual selection may have played a more important role in the distant origin and evolution of mammals earlier than previously thought. Sexual selection may explain the subsequent independent evolution of cranial outgrowths and pachyostosis in different therapsid lineages (Biarmosuchia, Dinocephalia, Gorgonopsia and Dicynodontia). [ABSTRACT FROM AUTHOR]

A skeleton discovered in the Upper Cretaceous Sierra Barrosa Formation (Turonian-Coniacian) of Neuquén Province, Argentina represents a new species of theropod dinosaur related to the long snouted, highly pneumatized Megaraptoridae. The holotype specimen of Murusraptor barrosaensis n.gen et n.sp. (MCF-PVPH-411) includes much of the skull, axial skeleton, pelvis and tibia. Murusraptor is unique in having several diagnostic features that include anterodorsal process of lacrimal longer than height of preorbital process, and a thick, shelf-like thickening on the lateral surface of surangular ventral to the groove between the anterior surangular foramen and the insert for the uppermost intramandibular process of the dentary. Other characteristic features of Murusraptor barrosaensis n.gen. et n. sp.include a large mandibular fenestra, distal ends of caudal neural spines laterally thickened into lateral knob-like processes, short ischia distally flattened and slightly expanded dorsoventrally. Murusraptor belongs to a Patagonian radiation of megaraptorids together with Aerosteon, Megaraptor and Orkoraptor. In spite being immature, it is a larger but more gracile animal than existing specimens of Megaraptor, and is comparable in size with Aerosteon and Orkoraptor. The controversial phylogeny of the Megaraptoridae as members of the Allosauroidea or a clade of Coelurosauria is considered analyzing two alternative data sets. [ABSTRACT FROM AUTHOR]

Although there are nine genera of ctenochasmatoids reported from the Jehol Biota, at present each is known from a specimen that has either a skull or a relatively complete postcranial skeleton. A nearly complete juvenile specimen of Gladocephaloideus from the Lower Cretaceous Jiufotang Formation of Sihedang, Lingyuan of Liaoning Province is the most complete ctenochasmatoid preserved to date with a skull and postcranial skeleton. Based on the holotype (IG-CAGS 08–07) and the nearly complete new specimen (JPM 2014–004), the diagnosis of Gladocephaloideus is amended: approximately 50 teeth in total with sharp tips; small nasoantorbital opening, occupying approximately 13% of total skull length; ratio of prenarial rostrum length to skull length approximately 0.63; deep groove along the mid-line of the mandibular symphysis; length to width ratio of the longest cervical vertebra = 4.1; ratio of femur length to tibia length = 0.61; tibia as long as the wing-phalange 1. Phylogenetic analysis recovers Gladocephaloideus within the clade Ctenochasmatidae. Gladocephaloideus has a closer relationship to the Chinese Pterofiltrus rather than to other ctenochasmatid pterosaurs. Microstructure of limb bones implies that JPM 2014–004 represents an early juvenile of Gladocephaloideus jingangshanensis, and that the type specimen is not a fully grown specimen either. We assume that the holotype may equate to the late juvenile or sub-adult developmental stage of Gladocephaloideus. [ABSTRACT FROM AUTHOR]

The Upper Cretaceous (middle-late Campanian) Wahweap Formation of southern Utah contains the oldest diagnostic evidence of ceratopsids (to date, all centrosaurines) in North America, with a number of specimens recovered from throughout a unit that spans between 81 and 77 Ma. Only a single specimen has been formally named, Diabloceratops eatoni, from the lower middle member of the formation. Machairoceratops cronusi gen. et sp. nov., a new centrosaurine ceratopsid from the upper member of the Wahweap Formation, is here described based on cranial material representing a single individual recovered from a calcareous mudstone. The specimen consists of two curved and elongate orbital horncores, a left jugal, a nearly complete, slightly deformed braincase, the left squamosal, and a mostly complete parietal ornamented by posteriorly projected, anterodorsally curved, elongate spikes on either side of a midline embayment. The fan-shaped, stepped-squamosal is diagnostic of Centrosaurinae, however, this element differs from the rectangular squamosal in Diabloceratops. Machairoceratops also differs in the possession of two anterodorsally (rather than laterally) curved epiparietal ornamentations on either side of a midline embayment that are distinguished by a posteromedially-oriented sulcus along the entire length of the epiparietal. Additionally, the parietosquamosal frill is lacking any other epiossifications along its periphery. Machairoceratops shares a triangular (rather than round) frill and spike-like epiparietal loci (p1) ornamentation with the stratigraphically lower Diabloceratops. Both parsimony and Bayesian phylogenetic analyses place Machairoceratops as an early-branching centrosaurine. However, the parsimony-based analysis provides little resolution for the position of the new taxon, placing it in an unresolved polytomy with Diabloceratops. The resultant Bayesian topology yielded better resolution, aligning Machairoceratops as the definitive sister taxon to a clade formed by Diabloceratops and Albertaceratops. Considered together, both phylogenetic methods unequivocally place Machairoceratops as an early-branching centrosaurine, and given the biostratigraphic position of Machairoceratops, these details increase the known ceratopsid diversity from both the Wahweap Formation and the southern portion of Laramidia. Finally, the unique morphology of the parietal ornamentation highlights the evolutionary disparity of frill ornamentation near the base of Centrosaurinae. [ABSTRACT FROM AUTHOR]

This study reports on a new ceratopsid, Spiclypeus shipporum gen et sp. nov., from the lower Coal Ridge Member of the Judith River Formation in Montana, USA, which dates to ~76 Ma (upper Campanian). The species is distinguished by rugose dorsal contacts on the premaxillae for the nasals, laterally projecting postorbital horncores, fully fused and anteriorly curled P1 and P2 epiparietals, and a posterodorsally projecting P3 epiparietal. The holotype specimen is also notable for its pathological left squamosal and humerus, which show varied signs of osteomyelitis and osteoarthritis. Although the postorbital horncores of Spiclypeus closely resemble those of the contemporaneous ‘Ceratops’, the horncores of both genera are nevertheless indistinguishable from those of some other horned dinosaurs, including Albertaceratops and Kosmoceratops; ‘Ceratops’ is therefore maintained as a nomen dubium. Cladistic analysis recovers Spiclypeus as the sister taxon to the clade Vagaceratops + Kosmoceratops, and appears transitional in the morphology of its epiparietals. The discovery of Spiclypeus adds to the poorly known dinosaur fauna of the Judith River Formation, and suggests faunal turnover within the formation. [ABSTRACT FROM AUTHOR]

We describe Sarmientosaurus musacchioi gen. et sp. nov., a titanosaurian sauropod dinosaur from the Upper Cretaceous (Cenomanian—Turonian) Lower Member of the Bajo Barreal Formation of southern Chubut Province in central Patagonia, Argentina. The holotypic and only known specimen consists of an articulated, virtually complete skull and part of the cranial and middle cervical series. Sarmientosaurus exhibits the following distinctive features that we interpret as autapomorphies: (1) maximum diameter of orbit nearly 40% rostrocaudal length of cranium; (2) complex maxilla—lacrimal articulation, in which the lacrimal clasps the ascending ramus of the maxilla; (3) medial edge of caudal sector of maxillary ascending ramus bordering bony nasal aperture with low but distinct ridge; (4) ‘tongue-like’ ventral process of quadratojugal that overlaps quadrate caudally; (5) separate foramina for all three branches of the trigeminal nerve; (6) absence of median venous canal connecting infundibular region to ventral part of brainstem; (7) subvertical premaxillary, procumbent maxillary, and recumbent dentary teeth; (8) cervical vertebrae with ‘strut-like’ centroprezygapophyseal laminae; (9) extremely elongate and slender ossified tendon positioned ventrolateral to cervical vertebrae and ribs. The cranial endocast of Sarmientosaurus preserves some of the most complete information obtained to date regarding the brain and sensory systems of sauropods. Phylogenetic analysis recovers the new taxon as a basal member of Lithostrotia, as the most plesiomorphic titanosaurian to be preserved with a complete skull. Sarmientosaurus provides a wealth of new cranial evidence that reaffirms the close relationship of titanosaurs to Brachiosauridae. Moreover, the presence of the relatively derived lithostrotian Tapuiasaurus in Aptian deposits indicates that the new Patagonian genus represents a ‘ghost lineage’ with a comparatively plesiomorphic craniodental form, the evolutionary history of which is missing for at least 13 million years of the Cretaceous. The skull anatomy of Sarmientosaurus suggests that multiple titanosaurian species with dissimilar cranial structures coexisted in the early Late Cretaceous of southern South America. Furthermore, the new taxon possesses a number of distinctive morphologies—such as the ossified cervical tendon, extremely pneumatized cervical vertebrae, and a habitually downward-facing snout—that have rarely, if ever, been documented in other titanosaurs, thus broadening our understanding of the anatomical diversity of this remarkable sauropod clade. The latter two features were convergently acquired by at least one penecontemporaneous diplodocoid, and may represent mutual specializations for consuming low-growing vegetation. [ABSTRACT FROM AUTHOR]

While centrosaurines and ceratopsids in general are abundant in the Late Campanian of northern Laramidia, they are much less commonly found in southern Laramidia. This has supported hypotheses of dinosaur provinciality and endemism in the Late Cretaceous with the delineation of at least two separate faunal zones, north and south Laramidia. There have been 12 genera of centrosaurines recognized from northern Laramidia while two genera, Diabloceratops and Nasutoceratops, have been named from southern Laramidia. We present an osteological description and taphonomic outline for a new centrosaurine ceratopsid from the Aguja Formation of northern Coahuila, Mexico that is not currently diagnosable to the generic level, but likely represents a new taxon. Further, we have included three-dimensional surface scans of all material attributed to this animal. Considering the large number of centrosaurines from northern Laramidia, it is likely that cladistic analyses are biased towards this faunal zone. New findings of southern centrosaurines are needed to correct this bias. This discovery expands the range of centrosaurines south to Coahuila, Mexico and adds new information to better characterize the morphology and taxonomy of centrosaurines from southern Laramidia and their evolution in comparison to their northern counterparts. [ABSTRACT FROM AUTHOR]

The jaw function of Smilodon fatalis has long been a source of debate. Although modern-day lions subdue large prey through the use of a suffocating throat bite, the dramatically elongated maxillary canines of S. fatalis suggest an alternative bite mechanism. The current literature favors a “canine shear-bite,” in which the depression of the cranium by the ventral neck flexors assists the mandibular adductors in closing the jaws. Although the model makes intuitive sense and appears to be supported by scientific data, the mechanical feasibility of “neck-powered” biting has not been experimentally demonstrated. In the present study, the computer-assisted manipulation of digitized images of a high-quality replica of an S. fatalis neck and skull shows that a rotation of the cranium by the ventral neck flexors will not result in jaw closure. Instead, the cranium and mandible rotate ventrally together (at the atlantooccipital joint), and the jaws remain in an open configuration. The only manner by which rotation of the cranium can simultaneously result in jaw closure is by an anterior rotation at the temporomandibular joint. Based on this finding, the author proposes a new Class 1 lever mechanism for S. fatalis jaw function. In this model, the mandible is immobilized against the neck of the prey and a dorsally directed force from the extension of the forelimbs rotates the cranium anteriorly at the temporomandibular joint. The maxillary canines pierce the prey’s neck and assist in clamping the ventral neck structures. The model is based on a maximum gape angle of approximately 90° and incorporates a secondary virtual point of rotation located slightly anteroventral to the temporomandibular joint. The Class 1 Lever Model is mechanically feasible, consistent with current data on S. fatalis anatomy and ecology, and may provide a basis for similar studies on other fossil taxa. [ABSTRACT FROM AUTHOR]

Pachycephalosaurids are small, herbivorous dinosaurs with domed skulls formed by massive thickening of the cranial roof. The function of the dome has been a focus of debate: the dome has variously been interpreted as the product of sexual selection, as an adaptation for species recognition, or as a weapon employed in intraspecific combat, where it was used in butting matches as in extant ungulates. This last hypothesis is supported by the recent identification of cranial pathologies in pachycephalosaurids, which appear to represent infections resulting from trauma. However, the frequency and distribution of pathologies have not been studied in a systematic fashion. Here, we show that pachycephalosaurids are characterized by a remarkably high incidence of cranial injury, where 22% of specimens have lesions on the dome. Frequency of injury shows no significant difference between different genera, but flat-headed morphs (here interpreted as juveniles or females) lack lesions. Mapping of injuries onto a digitial pachycephalosaurid skull shows that although lesions are distributed across the dome, they cluster near the apex, which is consistent with the hypothesis that the dome functioned for intraspecific butting matches. [ABSTRACT FROM AUTHOR]

Extinct animal behavior has often been inferred from qualitative assessments of relative brain region size in fossil endocranial casts. For instance, flight capability in pterosaurs and early birds has been inferred from the relative size of the cerebellar flocculus, which in life protrudes from the lateral surface of the cerebellum. A primary role of the flocculus is to integrate sensory information about head rotation and translation to stabilize visual gaze via the vestibulo-occular reflex (VOR). Because gaze stabilization is a critical aspect of flight, some authors have suggested that the flocculus is enlarged in flying species. Whether this can be further extended to a floccular expansion in highly maneuverable flying species or floccular reduction in flightless species is unknown. Here, we used micro computed-tomography to reconstruct “virtual” endocranial casts of 60 extant bird species, to extract the same level of anatomical information offered by fossils. Volumes of the floccular fossa and entire brain cavity were measured and these values correlated with four indices of flying behavior. Although a weak positive relationship was found between floccular fossa size and brachial index, no significant relationship was found between floccular fossa size and any other flight mode classification. These findings could be the result of the bony endocranium inaccurately reflecting the size of the neural flocculus, but might also reflect the importance of the flocculus for all modes of locomotion in birds. We therefore conclude that the relative size of the flocculus of endocranial casts is an unreliable predictor of locomotor behavior in extinct birds, and probably also pterosaurs and non-avian dinosaurs. [ABSTRACT FROM AUTHOR]