Cranial Anatomy and Palaeoneurology of the Archosaur Riojasuchus tenuisceps from the Los Colorados Formation,La Rioja,Argentina

Riojasuchus tenuisceps Bonaparte 1967 is currently known from four specimens, including two complete skulls, collected in the late 1960s from the upper levels of the Los Colorados Formation (Late Triassic), La Rioja, Argentina. Computed tomography (CT) scans of the skulls of the holotype and a referred specimen of Riojasuchus tenuisceps and the repreparation of the latter allows recognition of new features for a detailed analysis of its cranial anatomy and its comparison with a wide variety of other archosauriform taxa. The diagnosis of Riojasuchus tenuisceps is emended and two autapomorphies are identified on the skull: (1) a deep antorbital fossa with its anterior and ventral edges almost coinciding with the same edges of the maxilla itself and (2) a suborbital fenestra equal in size to the palatine-pterygoid fenestra. Also, the first digital 3D reconstruction of the encephalon of Riojasuchus tenuisceps was carried out to study its neuroanatomy, showing a shape and cranial nerve disposition consistent to that of other pseudosuchians.     

New observations on the skull of Archaeopteryx

Although skeletal remains of the iconic oldest known avialian Archaeopteryx have been known for almost 150 years, several aspects of the cranial anatomy of this taxon have remained enigmatic, mainly because of the strongly flattened and often fractured and incomplete nature of available skull materials. New investigation of the skulls of the recently described, excellently preserved tenth (Thermopolis) and the seventh (Munich) specimens revealed several previously unrecognized characters and helps to resolve some problematic issues. Thus, the nasal of Archaeopteryx shows a lateral notch for the lacrimal, as is found in many other saurischian dinosaurs, the maxilla clearly participates in the margin of the external nares, and there seems to be a pneumatic foramen in the lacrimal, comparable to the lacrimal fenestra found in many non-avian theropods. In the braincase, Archaeopteryx shows pneumatic features reminiscent of non-avian theropods, including a ventral basisphenoid recess and an anterior tympanic recess that is laterally incised into the basisphenoid/prootic. Most importantly, however, the postorbital process of the jugal shows a facet for the suture with the postorbital, thus resolving the question of whether Archaeopteryx had a closed postorbital bar. A new reconstruction of the skull of Archaeopteryx is presented, making the skull of this taxon even more theropod-like than previously recognized. Furthermore, the closed postorbital bar and the configuration of the bones of the skull roof cast serious doubt on claims that an avian-style cranial kinesis was present in this taxon.     

Lipiniella prima sp.nov. (Diptera,Chironomidae), larva and karyotype

The karyological and morphological features of the new speciesL. prima sp.nov. are described.L. prima has a 2n=8, three chromosome pairs are metacentric, the other is an acrocentric.L. prima has a high level of chromosomal polymorphism. There are distinct morphological features allowing to identify the larva ofL. prima.     

Cranial morphology of the Early Permian mesosaurid Mesosaurus tenuidens and the evolution of the lower temporal fenestration reassessed

The Early Permian mesosaurids are the oldest known aquatic amniotes with an exclusively Gondwanan distribution. Although several hundred of complete skeletons have been discovered and intensively studied, the anatomy and taxonomic composition of the group, as well as its phylogenetic relationships remain controversial. Several well-preserved mesosaurid specimens found in Uruguay justify a new anatomical reconstruction of the skull of Mesosaurus tenuidens, differing from earlier ones especially in the presence of a lower temporal fenestra. The significance of this structure for the evolution of temporal fenestration in amniotes is evaluated according to the two most recent phylogenetic hypotheses, in which mesosaurids are basalmost sauropsids or basalmost parareptiles. A synapsid-like fenestration may be the primitive condition for Amniota, and it may be also a basal condition for parareptiles, because recent phylogenies suggest a basal position for mesosaurids and lanthanosuchoids within that group, and both possess a lower temporal fenestra. Our results also give a moderately strengthened support for diapsid affinities of turtles.     

Ostéologie et affinités de Trematochampsa taqueti (Crocodylia,Mesosuchia) du Sénonien inférieur d'In Beceten (République du Niger)

Among the abundant remains of Mesosuchia that have been yielded by the early Senonian locality of In Beceten (Niger), the most numerous belong to the species Trematochampsa taquetiBuffetaut, 1974. The osteology of this species is described here. This medium-sized crocodilian, with a moderately elongated skull, is characterized, among other features, by its antorbital fenestra, its surangular-quadratojugal articulation, its slightly displaced post-orbital pillar, its teeth with wrinkled enamel, and its amphicoelous vertebrae. In its general outlook, this animal is rather reminiscent of the Goniopholidae, from the Upper Jurassic and the Cretaceous of Laurasia, but it is more primitive than them, and resemblances are probably due to convergence phenomena. The family Trematochampsidae is probably essentially Gondwanian, and could persist until the Senonian thanks to a certain isolation from Laurasia, where more progressive Crocodylia became predominant much earlier than in Africa and South America.     

A new galesaurid (Therapsida: Cynodontia) from the Lower Triassic of South Africa

A new galesaurid cynodont, Progalesaurus lootsbergensis gen. et sp. nov., is described on the basis of a well-preserved skull, lower jaw, right scapula and left atlantal neural arch. Autapomorphies of Progalesaurus include postcanine teeth bearing numerous mesial and distal accessory cusps that flank a recurved main cusp, a post-temporal fenestra bordered by the squamosal ventrally and a large external naris. Progalesaurus is similar to Galesaurus in possessing a poorly defined masseteric fossa on the dentary, a strongly recurved main cusp of the postcanine dentition, an incomplete secondary palate and a similar basisphenoid-parasphenoid morphology. A cladistic analysis of ten early cynodont genera resolves a monophyletic Galesauridae encompassing Cynosaurus , Progalesaurus and Galesaurus , although support for this clade is weak. Procynosuchus and Dvinia are placed at the base of Cynodontia whereas Thrinaxodon and Platycraniellus are positioned higher, but outside of Eucynodontia. The holotype and only known specimen of Progalesaurus was collected during systematic prospecting of Permo/Triassic boundary strata at New Lootsberg Pass, Graaff-Reinet District, South Africa. The discovery of Progalesaurus increases the number of valid Early Triassic cynodonts to four and sheds light on the tempo of early cynodont diversification after the end-Permian mass extinction.     

A new species of Theriosuchus (Atoposauridae, Crocodylomorpha) from the Late Jurassic (Kimmeridgian) of Guimarota, Portugal

A new species of an atoposaurid crocodilian, Theriosuchus guimarotae, is described from the Late Jurassic (Kimmeridgian) of Portugal. Theriosuchus guimarotae can be distinguished from other species of Theriosuchus by a lateral surface of squamosal bevelled ventrally; a rounded, caudally projecting and dorsally sculptured caudolateral corner of the squamosal; a premaxillomaxillary suture aligned caudomedially in dorsal aspect; a minimum space between the supratemporal foramina that comprises one third of the total width of the cranial table; a minimum width of the frontal between the orbits that comprises one third of the maximum width of the skull at the orbits; a dentition that comprises only pseudocaniniform and lanceolate-shaped teeth; the presence of an external mandibular fenestra and all vertebral bodies amphicoelous. Its osteology also sheds light on the diagnosis of Theriosuchus within Atoposauridae. The material additionally includes specimens representative of several ontogenetic stages, each of which is discussed here. With its Late Jurassic age, T. guimarotae represents the oldest well-preserved material of Theriosuchus and reveals further knowledge about the palaeobiogeography of the genus in western Europe.     

CRANIAL CREST DEVELOPMENT IN THE AZHDARCHOID PTEROSAUR TUPUXUARA, WITH A REVIEW OF THE GENUS AND TAPEJARID MONOPHYLY

Abstract: A portion of pterosaur skull from the Romualdo Member of the Santana Formation (?Albian–?Turonian, Cretaceous) of north‐east Brazil provides new data on the morphology and ontogeny of azhdarchoid pterosaur cranial crests. The specimen consists of parts of the cranial bones posterodorsal to the nasoantorbital fenestra, including partial nasals, lacrimals, frontals and possibly the parietals. A posterodorsally directed premaxillary crest with a concave posterior border is located dorsal to the posterior border of the nasoantorbital fenestra. A well‐defined suture indicates overlapping, posterodorsally directed growth of the premaxilla over the skull roof, suggesting that the generation of the premaxillary crest is a late ontogenetic feature and thus probably related to sexual display. The systematics of Tupuxuara and its relationship to other azhdarchoids is reviewed and a cladistic analysis of the group is presented. Tupuxuara is found to be the sister‐taxon to Azhdarchidae. Tupuxuara longicristatus Kellner and Campos, 1988 is argued to be the only valid named species in this genus and Thalassodromeus Kellner and Campos, 2002 is considered a junior subjective synonym of this taxon. As originally conceived, Tapejaridae is paraphyletic: a new, more restrictive version of Tapejaridae (including Tapejara and Sinopterus dongi) might exist, but its monophyly is weakly supported. Furthermore, Tapejara was found to be paraphyletic in all trees.     

New information on body size and cranial display structures of Pterodactylus antiquus, with a revision of the genus

Pterodactylus antiquus has long been thought to have been quite small (~50 cm wingspan) and to have differed from P. longicollum, Ctenochasma, Germanodactylus, and Gnathosaurus in lacking a bony cranial crest, though a soft tissue crest and occipital lappet have been described. This article describes a new specimen of P. antiquus larger than all previously known specimens, which demonstrates that the species exceeded 1 m in wingspan and had a low bony cranial crest. A smaller, incipient crest was identified on the holotype specimen. Additional specimens, including the counterpart of Wellnhofer’s original occipital lappet specimen, provide evidence of the occipital lappet and the soft tissue crest extending upward above the naso-antorbital fenestra and orbit. In order to provide a proper taxonomic context for the findings, the recent synonymization of the species Pterodactylus antiquus and P. kochi on the basis of shared correlation of tooth number and skull length despite perceived differences in dentition and skull, neck, and trunk proportions is reviewed. A measurement error that had made it appear that P. antiquus differed significantly from P. kochi in proportions is documented, and after correction of the measurement error and reevaluation of the dental evidence there are no significant differences between the two nominal species. Thus, the synonymization of P. antiquus and P. kochi was appropriate, and a revised diagnosis is presented. In addition, the species P. longicollum and P. micronyx, which for some years have been viewed as not congeneric with P. antiquus, are placed in a new genus and transferred to Aurorazhdarcho, respectively.     

The morphology and post-hatching development of the skull of Bolitoglossa nicefori (Caudata: Plethodontidae): developmental implications of recapitulation and repatterning

The cranial morphology of the direct-developing salamander Bolitoglossa nicefori and its post-hatching development are described and compared with that of other urodeles. Four stages of cranial development are defined on the basis of conspicuous events that occur during post-hatching ontogeny. The adult skull morphology of B. nicefori is similar to that of other plethodontids; however, some regions show interspecific variation. The post-hatching ontogeny of the skull and the stage of ossification observed in the hatchlings of B. nicefori show two important ontogenetic features: (1) a mosaic of early larval, metamorphic and post-metamorphic skull features in hatchlings, and (2) absence of characteristic larval elements in skull and hyoid apparatus. The distinctive stage of ossification in the hatchlings of B. nicefori could be caused by heterochronic changes in the ossification sequence, compared to the ontogeny of metamorphic salamanders. The possible heterochronic changes and the absence of larval traits are perhaps due to ontogenetic repatterning, yet without an obvious impact on the adult skull morphology (absence of morphological novelties). This might indicate a compartmentalized development. Further studies should be performed in order to establish the possible occurrence of recapitulatory patterns or ontogenetic repatterning in the skull morphogenesis of B. nicefori during its embryonic development.     

Comparative Anatomy of the Petrosal Bone of Dichobunoids,Early Members of Artiodactylamorpha (Mammalia)

Among Artiodactylamorpha, dichobunoids are some of the oldest fossil species that have been associated with Artiodactyla, the crown clade that includes hippopotamids, camelids, suoids, ruminants, and cetaceans. These important fossil species are known from early Eocene rocks of North America, Europe, and Asia, but their phylogenetic position has yet to be well resolved. Before generating such a phylogeny, it is first critical to document all of the anatomy of known dichobunoid fossils. Here we use CT scans to describe previously undescribed anatomy of the petrosal bone, a complex part of the mammalian skull that contains many variable and phylogenetically informative features. Results show that these extinct species share a number of features that are not documented in modern species including a lateral process of the epitympanic wing constituting the medial border of the piriform fenestra, and a tegmen tympani foramen that may have given passage to the ramus superior of the stapedial artery. Future comprehensive phylogenetic studies may show that many of these characters are plesiomophic for Artiodactylamopha. Some species (Diacodexis, Homacodon and ?Helohyus) exhibit a dorsolateral exposure of the mastoid region of the petrosal on the temporal part of the cranium. This uncommon feature has, to our knowledge, not been reported in another euungulate group.     

A New Baurusuchid (Crocodyliformes, Mesoeucrocodylia) from the Late Cretaceous of Brazil and the Phylogeny of Baurusuchidae

Background

Baurusuchidae is a group of extinct Crocodyliformes with peculiar, dog-faced skulls, hypertrophied canines, and terrestrial, cursorial limb morphologies. Their importance for crocodyliform evolution and biogeography is widely recognized, and many new taxa have been recently described. In most phylogenetic analyses of Mesoeucrocodylia, the entire clade is represented only by Baurusuchus pachecoi, and no work has attempted to study the internal relationships of the group or diagnose the clade and its members.

Methodology/Principal Findings

Based on a nearly complete skull and a referred partial skull and lower jaw, we describe a new baurusuchid from the Vale do Rio do Peixe Formation (Bauru Group), Late Cretaceous of Brazil. The taxon is diagnosed by a suite of characters that include: four maxillary teeth, supratemporal fenestra with equally developed medial and anterior rims, four laterally visible quadrate fenestrae, lateral Eustachian foramina larger than medial Eustachian foramen, deep depression on the dorsal surface of pterygoid wing. The new taxon was compared to all other baurusuchids and their internal relationships were examined based on the maximum parsimony analysis of a discrete morphological data matrix.

Conclusion

The monophyly of Baurusuchidae is supported by a large number of unique characters implying an equally large morphological gap between the clade and its immediate outgroups. A complex phylogeny of baurusuchids was recovered. The internal branch pattern suggests two main lineages, one with a relatively broad geographical range between Argentina and Brazil (Pissarrachampsinae), which includes the new taxon, and an endemic clade of the Bauru Group in Brazil (Baurusuchinae).     

Morphology of the interorbital region of Saimiri sciureus

The skull of the platyrrhine primate Saimiri sciureus is distinguished by a large interorbital fenestra. Juvenile skulls still show a bony interorbital septum with some small gaps. A morphogenetic study was undertaken to better understand the structures of the interorbital region, which represents a linkage between the base of the braincase and the nasal skeleton. Already in early ontogenetic stages a reduction of the posterior portion of the nasal capsule and of the cartilaginous interorbital septum are observed, resulting in the formation of a primary interorbital fenestra. A bony interorbital septum is mainly formed in perinatal age stages by ossification of the presphenoid and by medial fusion of the frontals; the primary interorbital fenestra is retained as a small opening. It only occurs in late juvenile stages when the definitive interorbital fenestra develops by by secondary transformation of bone into a membrane of dense connective tissue; this process is most probably caused by mechanical friction of the very closely approximated eyes of both sides.     

A Second Soundly Sleeping Dragon: New Anatomical Details of the Chinese Troodontid Mei long with Implications for Phylogeny and Taphonomy

A second nearly complete, articulated specimen of the basal troodontid Mei long (DNHM D2154) is reported from the Early Cretaceous (Hauterivian-Valanginian) lower Yixian Formation, Liaoning Province, China. New diagnostic features of Mei long are identified, including: a uniquely shaped maxilla, low with small, low maxillary fenestra; sacrum with an extremely wide caudal portion and elongate 4^th and 5^th sacral processes; and a large distal articular surface on the tibiotarsus which continues caudally on the tibia. A phylogenetic analysis including new data from the second specimen recovered Mei as a basal troodontid, in keeping with previous analyses. Although the skeleton exhibits several juvenile-like features including free cervical ribs, unfused frontals and nasals, and a short snouted skull, other attributes, full fusion of all neurocentral synostoses and the sacrum, and dense exteriors to cortical bone, suggest a small, mature individual. Microscopic examination of tibia and fibula histology confirms maturity and suggests an individual greater than two years old with slowed growth. Despite being one of the smallest dinosaurs, Mei long exhibits multi-year growth and cortical bone consisting largely of fibro-lamellar tissue marked by lines of arrested growth as in much larger and more basal theropods. This Mei long specimen lies in a similar but mirrored sleeping position to that of the holotype, strengthening the hypothesis that both specimens were preserved in a stereotypical life position. Like many Liaoning specimens, the new specimen also lacks extensive taphonomic and stratigraphic data, making further behavioral inference problematic.     

A New Species of Garjainia Ochev, 1958 (Diapsida: Archosauriformes: Erythrosuchidae) from the Early Triassic of South Africa

A new species of the erythrosuchid archosauriform reptile Garjainia Ochev, 1958 is described on the basis of disarticulated but abundant and well-preserved cranial and postcranial material from the late Early Triassic (late Olenekian) Subzone A of the Cynognathus Assemblage Zone of the Burgersdorp Formation (Beaufort Group) of the Karoo Basin of South Africa. The new species, G. madiba, differs from its unique congener, G. prima from the late Olenekian of European Russia, most notably in having large bony bosses on the lateral surfaces of the jugals and postorbitals. The new species also has more teeth and a proportionately longer postacetabular process of the ilium than G. prima. Analysis of G. madiba bone histology reveals thick compact cortices comprised of highly vascularized, rapidly forming fibro-lamellar bone tissue, similar to Erythrosuchus africanus from Subzone B of the Cynognathus Assemblage Zone. The most notable differences between the two taxa are the predominance of a radiating vascular network and presence of annuli in the limb bones of G. madiba. These features indicate rapid growth rates, consistent with data for many other Triassic archosauriforms, but also a high degree of developmental plasticity as growth remained flexible. The diagnoses of Garjainia and of Erythrosuchidae are addressed and revised. Garjainia madiba is the geologically oldest erythrosuchid known from the Southern Hemisphere, and demonstrates that erythrosuchids achieved a cosmopolitan biogeographical distribution by the end of the Early Triassic, within five million years of the end-Permian mass extinction event. It provides new insights into the diversity of the Subzone A vertebrate assemblage, which partially fills a major gap between classic ‘faunal’ assemblages from the older Lystrosaurus Assemblage Zone (earliest Triassic) and the younger Subzone B of the Cynognathus Assemblage Zone (early Middle Triassic).     

Skull of Grippia Longirostris: no contradiction with a diapsid affinity for the Ichthyopterygia

The skull of Grippia longirostris , one of the basal ichthyopterygians, is redescribed, based on a newly prepared referred specimen and the examination of others. Grippia had been considered to be the most basal ichthyopterygian until recently, and it has been suggested that the quadratojugal enters the upper temporal fenestra in this species. This and other unique features, which were considered plesiomorphic for ichthyopterygians, obscured the relationship of the group to other amniotes. It is shown here that the quadratojugal is much smaller than previously described, and does not enter the upper temporal fenestra. The skull configuration of the species is very similar to the condition in the recently-described skull of Utatsusaurus hataii , another ichthyopterygian belonging to the basal grade of the group. The close similarity between the two help clarify ichthyopterygian plesiomorphies. There is no feature in the skull of Grippia that particularly contradicts the emerging consensus that ichthyopterygians are diapsids.     

Diverse radial symmetry among the Cambrian Fortunian fossil embryos from northern Sichuan and southern Shaanxi provinces,South China

The Cambrian Fortunian fossil embryos exhibit embryonic development of ancient animals and hence have important bearings on evolutionary developmental biology. They have radial symmetry, and may be early representatives of cnidarians. Here we report new material of three-dimensionally phosphatized fossil embryos from the Fortunian Kuanchuanpu Formation and coeval strata in northern Sichuan and southern Shaanxi provinces, South China. The new material includes previously reported fossil embryos assignable to Pseudooides prima with biradial symmetry or pseudo-hexaradial symmetry, Quadrapyrgites quadratacris with tetraradial symmetry, and Olivooides multisulcatus with pentaradial symmetry. Additionally, we recovered two new types of fossil embryos, i.e., Embryo I with hexaradial symmetry and Embryo II with octaradial symmetry, and they are tentatively suggested to represent new cnidarians. In contrast to the diverse radial symmetry of the Fortunian cnidarians, modern cnidarians exhibit stable tetraradial symmetry in medusozoans, biradial symmetry in anthozoans, and bilateral symmetry in siphonophores (Hydrozoa). The current study supports the view that the tetraradial symmetry of modern medusozoans is a surviving remnant of their Fortunian relatives.