Antorbital Fenestra Research Articles

Cranial anatomy of Bagualia alba (Dinosauria, Eusauropoda) from the Early Jurassic of Patagonia and the implications for sauropod cranial evolution

Sauropods are the largest terrestrial vertebrates ever known. During their evolution, they underwent numerous morphological changes, some of which occurred at the end of the Early Jurassic. However, current knowledge of Early Jurassic dinosaurs is scarce, especially cranial information. Here we describe the skull of the basal eusauropod Bagualia alba from the Cañadón Asfalto Formation (Toarcian; Early Jurassic) of Patagonia, Argentina, which represents the most complete eusauropod skull from the late Early Jurassic worldwide. The osteological description and phylogenetic relationships of Bagualia allowed us to determine the acquisition of the common cranial bauplan of eusauropods at the end of the Early Jurassic: a tall and robust rostrum with great retraction of the external nares; a deep narial fossa; a reduced antorbital fenestra; verticalization of the lacrimal; prefrontal without an anterior process; anteroposterior shortening of the frontal; a supratemporal fenestra visible laterally; a robust braincase; ‘U’-shaped jaws in dorsal view; reduction in the number of teeth in the dentary; teeth with crown to crown occlusion; development of lateral plates in the premaxilla and maxilla; and a dorsoventral expansion of the dentary symphysis. These cranial characteristics were recently suggested to be related to a change in the diet of the sauropods toward mainly hard vegetation. In this context, Bagualia contributes to filling a gap in cranial evolution between the gracile skulls of Sauropodiformes and the highly robust skulls widely represented in Middle Jurassic taxa.

Cranial osteology and paleoneurology of Tarjadia ruthae: An erpetosuchid pseudosuchian from the Triassic Chañares Formation (late Ladinian-?early Carnian) of Argentina.

Tarjadia ruthae is a quadrupedal terrestrial pseudosuchian from the Middle-early Upper Triassic of the Chañares Formation, La Rioja Province, Argentina. Originally, this species was identified as an indeterminate archosaur and later as a doswelliid archosauriform based on very fragmentary specimens characterized by the ornamentation of the skull roof and osteoderms. Additional specimens (including skulls and postcrania) recovered in the last decade show that Tarjadia is an erpetosuchid, an enigmatic pseudosuchian group composed of six species registered in Middle-Upper Triassic continental units of Tanzania, Germany, Scotland, North America, Brazil, and Argentina. Tarjadia ruthae from Argentina and Parringtonia gracilis from Tanzania are the best preserved and more abundant species. Although the monophyly of Erpetosuchidae is well supported, alternative high-level positions within Archosauria have been suggested, such as sister taxon to Crocodylomorpha, Aetosauria, or Ornithosuchidae. In order to improve the knowledge about the erpetosuchids, we performed a detailed description and paleoneurological reconstruction of the skull of Tarjadia ruthae, based on two articulated partial skulls (CRILAR-Pv 478 and CRILAR-Pv 495) and other fragmentary specimens. We analyzed the stratigraphic and geographic occurrence of historical and new specimens of Tarjadia and provided a new emended diagnosis (the same for the genus as for the species, due to monotypy) along with a comparative description of the cranial endocast. The skull of Tarjadia is robust, with a thick and strongly ornamented skull roof, triangular in dorsal view, with concave lateral margins at mid-length that form an abrupt widened posterior region. The external nares are the smallest openings of the skull. The antorbital fossa is deeply excavated and has a small heart-shaped fenestra with both lobes pointing anteriorly. The supratemporal fenestrae are as large and rounded as the orbits, and the infratemporal fenestrae are L-shaped with an extensive excavation along the jugal, quadratojugal and quadrate. The hemimandibles are low, slightly concave on the dentigerous region and strongly convex on the posterior region, conferring them a S-shaped profile in dorsal view. The external mandibular fenestra is small and elliptic, being twice longer than high. The maxillary dentition is restricted to the anterior to mid region of the rostrum. Since the braincase of both specimens is partially damaged, the dorsal surface of the brain could not be entirely reconstructed. As a result, the endocast is anteroposteriorly elongated and seemingly flat, and the cephalic flexure seems to be lower than expected for a suchian. The labyrinth is twice wider than high, the semicircular canals are remarkably straight, and the anterior canal is longer than the posterior one.

A new spinosaurid dinosaur species from the Early Cretaceous of Cinctorres (Spain)

A new spinosaurid genus and species is described based on the right maxilla and five caudal vertebrae of a single specimen from the Arcillas de Morella Formation (Early Cretaceous) at the locality of Cinctorres (Castellón, Spain). Protathlitis cinctorrensis gen. et sp. nov. is diagnosed by one autapomorphic feature as well as by a unique combination of characters. The autapomorphy includes a subcircular depression in the anterior corner of the antorbital fossa in the maxilla. The new Iberian species is recovered as a basal baryonychine. The recognition of Protathlitis cinctorrensis gen. et sp. nov. as the first baryonychine dinosaur species identified from the Arcillas de Morella Formation (late Barremian) from the same time as Vallibonavenatrix cani, the first spinosaurine dinosaur from the same formation in the Morella subbasin (Maestrat Basin, eastern Spain), indicates that the Iberian Peninsula was home to a highly diverse assemblage of medium-to-large bodied spinosaurid dinosaurs. It seems that spinosaurids appeared during the Early Cretaceous in Laurasia, with the two subfamilies occupying the western part of Europe during this period. Later, during the Barremian–Aptian, they migrated to Africa and Asia, where they would diversify. In Europe, baryonychines were dominant, while in Africa, spinosaurines were most abundant.

A New Specimen of Sinosaurus triassicus (Dinosauria: Theropoda) from the Early Jurassic of Lufeng, Yunnan, China

ABSTRACT The holotype of Sinosaurus triassicus is poorly preserved, and no previous research has focused exclusively on its anatomy. This research reports a new specimen recovered near the locality of the holotype of S. triassicus including a complete skull with mandible and 11 cervical vertebrae. Three newly identified traits are proposed to be unique of the species – (1) a vertical crest developed dorsoventrally from the ventral portion of the maxilla to the nasal crest; (2) ventral border of the antorbital fenestra formed mostly by the anterior process of the jugal; and (3) a fenestra between the nasal, the lacrimal and the prefrontal. This research provides new information on the cranial anatomy of this species as well as the genus Sinosaurus, offering more accessibility for later research concerning the evolution of early deriving neotheropods.

Anatomy of the neornithischian dinosaur Parksosaurus warreni (Parks, 1926) from the Upper Cretaceous (lower Maastrichtian) Horseshoe Canyon Formation of Alberta, Canada

The fossil record of Late Cretaceous non-hadrosaurid neornithischians from the Western Interior of North America is sparse. Parksosaurus warreni is an early Maastrichtian taxon currently known only from a skull and much of the associated postcranial skeleton and a referred isolated tooth from the Horseshoe Canyon Formation of Alberta, Canada. This article presents a detailed anatomical account of the holotype of this species following extensive additional preparation of the holotype and provides photographs of the postcranial elements for the first time. The premaxilla has a dorsoventrally deep posterolateral process, which overlaps the maxilla. The antorbital fenestra is small and teardrop-shaped. The lingual surfaces of dentary teeth bear nine or 10 vertically extending ridges that terminate in apical denticles. The suprascapula is mineralized or ossified. The dorsal margin of the ischium is posterodorsally concave at mid-length. The greater and lesser trochanters of the femur are not separated by a distinct cleft. The pendent fourth trochanter is placed on the proximal half of the femur. The more distal caudal vertebrae have greatly elongated prezygapophyses, which, together with a basket of ossified tendons ensheathing the caudal vertebrae, indicate that the tail may have functioned as a dynamic stabilizer during locomotion. The substantial new anatomical information from the holotype and only known skeleton of Parksosaurus warreni will allow more detailed comparisons to other closely related taxa and facilitate further discussions of unresolved aspects of the interrelationships of neornithischian dinosaurs.

A new juvenile Yamaceratops (Dinosauria, Ceratopsia) from the Javkhlant Formation (Upper Cretaceous) of Mongolia.

Here we report a new articulated skeleton of Yamaceratops dorngobiensis (MPC-D 100/553) from the Khugenetjavkhlant locality at the Shine Us Khudag (Javkhlant Formation, ?Santonian-Campanian) of the eastern Gobi Desert, Mongolia, which represents the first substantially complete skeleton and the first juvenile individual of this taxon. The specimen includes a nearly complete cranium and large portions of the vertebral column and appendicular skeleton. Its skull is about 2/3 the size of the holotype specimen, based on mandibular length. Its juvenile ontogenetic stage is confirmed by multiple indicators of skeletal and morphological immaturity known in ceratopsians, such as the long-grained surface texture on the long bones, the smooth external surface on the postorbital, open neurocentral sutures of all caudal vertebrae, a large orbit relative to the postorbital and jugal, the low angle of the lacrimal ventral ramus relative to the maxillary teeth row, narrow frontal, and straight ventral edge of the dentary. Osteohistological analysis of MPC-D 100/553 recovered three lines of arrested growth, implying around 3 years of age when it died, and verified this specimen’s immature ontogenetic stage. The specimen adds a new autapomorphy of Yamaceratops, the anteroventral margin of the fungiform dorsal end of the lacrimal being excluded from the antorbital fossa. Furthermore, it shows a unique combination of diagnostic features of some other basal neoceratopsians: the ventrally hooked rostral bone as in Aquilops americanus and very tall middle caudal neural spines about or more than four times as high as the centrum as in Koreaceratops hwaseongensis, Montanoceratops cerorhynchus, and Protoceratops andrewsi. The jugal with the subtemporal ramus deeper than the suborbital ramus as in the holotype specimen is also shared with A. americanus, Liaoceratops yanzigouensis, and juvenile P. andrewsi. Adding 38 new scorings into the recent comprehensive data matrix of basal Neoceratopsia and taking into account the ontogenetically variable characters recovered Y. dorngobiensis as the sister taxon to Euceratopsia (Leptoceratopsidae plus Coronosauria). A second phylogenetic analysis with another matrix for Ceratopsia also supported this position. The new phylogenetic position of Y. dorngobiensis is important in ceratopsian evolution, as this taxon represents one of the basalmost neoceratopsians with a broad, thin frill and hyper-elongated middle caudal neural spines while still being bipedal.

THE PRESENCE OF AN ORBITOANTORBITAL FENESTRA: FURTHER EVIDENCE OF THE ANUROGNATHID PECULIARITY WITHIN THE PTEROSAURIA

The anurognathids are peculiar pterosaurs characterized by broad skulls with very short rostra and broadly arched jaws. The presence of distinct or confluent external naris and antorbital fenestra in these pterosaurs has been debated in the last years. The relatively well-preserved specimens of Batrachognathus volans show that the antorbital fenestra was confluent with the orbit forming an enormous orbitoantorbital fenestra. This feature is evident also in Jeholopterus ningchengensis. The consequent modification of the matrices of two recently published phylogenetic analyses about the in-group pterosaur relationships shows that the Anurognathidae are a derived clade of non-monofenestratan pterosaurs. Anurognathidae (including also 'Dimorphodon' weintraubi according to the definition by Hone 2020) are still a scarcely known clade because only a few specimens have been adequately described in the literature.

A new, remarkably preserved, enantiornithine bird from the Upper Cretaceous Qiupa Formation of Henan (central China) and convergent evolution between enantiornithines and modern birds

AbstractA new enantiornithine bird is described on the basis of a well preserved partial skeleton from the Upper Cretaceous Qiupa Formation of Henan Province (central China). It provides new evidence about the osteology of Late Cretaceous enantiornithines, which are mainly known from isolated bones; in contrast, Early Cretaceous forms are often represented by complete skeletons. While the postcranial skeleton shows the usual distinctive characters of enantiornithines, the skull displays several features, including confluence of the antorbital fenestra and the orbit and loss of the postorbital, evolved convergently with modern birds. Although some enantiornithines retained primitive cranial morphologies into the latest Cretaceous Period, at least one lineage evolved cranial modifications that parallel those in modern birds.

Paranasal sinus system and upper respiratory tract evolution in Mesozoic pelagic crocodylomorphs.

Thalattosuchians were a predominately marine clade of Mesozoic crocodylomorphs, including semi-aquatic teleosauroid and obligately pelagic metriorhynchid subclades. Recent advances in our understanding of thalattosuchian endocranial anatomy have revealed new details of the evolutionary transition from terrestrial to marine to pelagic taxa. Paranasal sinuses, however, have received little attention. Herein, we investigate the evolution of the paranasal sinus system and part of the upper respiratory system (nasopharyngeal ducts) in Thalattosuchia, by reconstructing the nasal and paranasal anatomy in CT scans of seven thalattosuchian skulls: one teleosauroid, two basal metriorhynchoids and four metriorhynchids. Our outgroups were: three extant crocodylian species (including adult and subadult skulls) and the basal crocodyliform Protosuchus. We found thalattosuchians exhibit exceptionally reduced paranasal sinus systems, solely comprising the antorbital sinus, as has been previously proposed. The semi-aquatic basal thalattosuchians Palgiopthalmosuchus gracilirostris and Pelagosaurus typus both have an antorbital sinus partially located medial to a reduced external antorbital fenestra and broadly communicating with the dorsal alveolar canal. In pelagic metriorhynchids, the antorbital cavity is more extensive than in basal taxa and possibly had an active function associated with a hypothesized accessory suborbital diverticulum, but our reconstructions are insufficient to confirm or reject the presence of such a diverticulum. The nasopharyngeal ducts of metriorhynchids are dorsoventrally enlarged, possibly enabling stronger ventilation. The sequence of acquisition of craniofacial adaptations show a mosaic pattern and appears to predate many skeletal adaptations, suggesting these changes occurred early in the thalattosuchian marine transition.

A neoceratopsian dinosaur from the early Cretaceous of Mongolia and the early evolution of ceratopsia

Ceratopsia is a diverse dinosaur clade from the Middle Jurassic to Late Cretaceous with early diversification in East Asia. However, the phylogeny of basal ceratopsians remains unclear. Here we report a new basal neoceratopsian dinosaur Beg tse based on a partial skull from Baruunbayan, Ömnögovi aimag, Mongolia. Beg is diagnosed by a unique combination of primitive and derived characters including a primitively deep premaxilla with four premaxillary teeth, a trapezoidal antorbital fossa with a poorly delineated anterior margin, very short dentary with an expanded and shallow groove on lateral surface, the derived presence of a robust jugal having a foramen on its anteromedial surface, and five equally spaced tubercles on the lateral ridge of the surangular. This is to our knowledge the earliest known occurrence of basal neoceratopsian in Mongolia, where this group was previously only known from Late Cretaceous strata. Phylogenetic analysis indicates that it is sister to all other neoceratopsian dinosaurs.

Dromaeosaurid crania demonstrate the progressive loss of facial pneumaticity in coelurosaurian dinosaurs

Abstract Dinosaurs are notable for their extensive skeletal pneumaticity, a feature that may have helped facilitate the development of various ‘extreme’ body plans in this group. Despite its relevance to understanding the evolution of the avian body plan, this feature has only been described in detail for a few non-avian dinosaurs, and cranial pneumaticity outside the braincase remains poorly documented. I describe facial pneumatic features in members of the Dromaeosauridae, a clade of hypercarnivorous dinosaurs closely allied to birds. Variation in the pneumaticity of the nasals and jugals, the position and shape of the pneumatic fenestrae of the maxilla and the border of the antorbital fossa shows that facial pneumaticity differed substantially among closely related dromaeosaurids and other bird-like dinosaurs. Ancestral state reconstructions of facial pneumaticity in coelurosaurs suggest a complex evolutionary history for these features. Surprisingly, the general trend along the path towards birds was the loss or reduction of superficial pneumatic features on the snout and cheek. Some facial pneumatic features seem to have evolved secondarily in some derived bird-like forms. The results show superficial facial pneumaticity did not increase in coelurosaurs and emphasize the complexity of the evolution of pneumatization in the lineage leading to birds.

A new specimen of Teleidosaurus calvadosii (Eudes-Deslongchamps, 1866) (Crocodylia, Thalattosuchia) from the Middle Jurassic of France

Our understanding of early metriorhynchid evolution, the highly specialised pelagic clade of Mesozoic crocodylomorphs, is hampered by the paucity of basal metriorhynchoids and how poorly preserved most of these fossils are. This means the transition from a ‘gharial-like’ body-plan to a ‘pseudo-dolphin-like’ one, which characterises metriorhynchoid evolution, is still poorly understood. Herein the Middle Jurassic taxon Teleidosaurus calvadosii (J.A. Eudes-Deslongchamps, 1866) is re-evaluated. The holotype of T. calvadosii was destroyed during World War Two, as were all referred specimens. All that survived are plaster casts of the holotype. However, a recently discovered specimen from the middle Bathonian of Ecouché, Normandy, France is herein referred to T. calvadosii. This specimen preserves the antorbital region, an incomplete mandible and one cervical vertebra. This specimen is designated as neotype of T. calvadosii. The new specimen allows to re-evaluate the anatomy of T. calvadosii, in particular the antorbital fenestra, which has been misunderstood. Moreover, a comparison is made between the French Middle Jurassic metriorhynchoids Teleidosaurus and Eoneustes. Both genera appear as valid and E. bathonicus is considered to be a nomen dubium.

Notes on the cheek region of the Late Jurassic theropod dinosaur Allosaurus.

Allosaurus, from the Late Jurassic of North America and Europe, is a model taxon for Jurassic basal tetanuran theropod dinosaurs. It has achieved an almost iconic status due to its early discovery in the late, 19th century, and due to the abundance of material from the Morrison Formation of the western U.S.A., making Allosaurus one of the best-known theropod taxa. Despite this, various aspects of the cranial anatomy of Allosaurus are surprisingly poorly understood. Here, we discuss the osteology of the cheek region, comprised by the jugal, maxilla, and lacrimal. This region of the skull is of importance for Allosaurus taxonomy and phylogeny, particularly because Allosaurus has traditionally been reconstructed with an unusual cheek configuration, and because the European species Allosaurus europaeus has been said to be different from North American material in the configuration of these bones. Based on re-examination of articulated and disarticulated material from a number of repositories, we show that the jugal participates in the antorbital fenestra, contradicting the common interpretation. The jugal laterally overlies the lacrimal, and forms an extended antorbital fossa with this bone. Furthermore, we document previously unrecorded pneumatic features of the jugal of Allosaurus.

Cranial morphology of a new phytosaur (Diapsida, Archosauria) from the Upper Triassic of India: implications for phytosaur phylogeny and biostratigraphy

AbstractDetailed description and phylogenetic assessment of a phytosaur skull collected from the Tiki Formation of the Rewa Gondwana Basin of India and earlier diagnosed asParasuchus hislopi, show that it pertains to a new genus and species,Volcanosuchus statisticae. The new taxon is characterized by marginal overlapping of the nostrils by the antorbital fenestrae, external nares situated on a bulbous and raised dome, the lateral surface of the jugal ornamented by a prominent ridge defined by multiple tubercles and radiating thread‐like structures, and distinct ornamentation patterns on the rostrum and skull table. Phylogenetic analysis nestsVolcanosuchuswithin Mystriosuchinae, where it forms a sister taxon to (Rutiodon + Leptosuchomorpha) and marks the transition between the basal Parasuchidae and more derived Mystriosuchinae phytosaurs. Evolution of the phytosaur skulls resulted in changes from non‐overlapping nostril and antorbital fenestra to an overlapping state, anteroposterior elongation of the exoccipital–supraoccipital shelf, appearance of a median ridge on the basioccipital, and reduction of the supratemporal fenestra. Considerable faunal overlap of the Tiki Formation is evident with the lower Maleri Formation, which is late Carnian based on the occurrence ofHyperodapedon,ParasuchusandExaeretodon. The Tiki Formation correlates with the Ischigualasto Formation of Argentina, the upper part of the Santa Maria Formation, and the overlying lower Caturrita Formation of Brazil, the Isalo II Beds of Madagascar, Lossiemouth Sandstone of Scotland, and the lower Tecovas Formation of the Chinle Group of North America, and ranges from late Carnian to early/middle Norian.

Scelidosaurus harrisonii from the Early Jurassic of Dorset, England: cranial anatomy

AbstractScelidosaurus harrisonii is an early (Late Sinemurian) armoured ornithischian dinosaur whose remains have, to date, only been recovered from a restricted location on the south coast of Dorset (Charmouth), England. This dinosaur has been known since 1859, but only on the basis of a partial description found in two articles published in the early 1860s by Richard Owen. The original material, discovered in 1858, comprised the majority of the skull and its associated postcranial skeleton, and represents the first ever, more or less complete dinosaur discovered. In addition to the original material, a number of further discoveries have been made at Charmouth; these latter supplement the information that can be gleaned from the original specimen. This article describes the skull of Scelidosaurus.The external surface of individual skull bones in ontogenetically relatively mature individuals displays exostoses, a patina of fibrous or granular-textured bone that anchored an external shielding of keratinous scales. There is a small, edentulous rostral beak, behind which is found a row of five heterodont premaxillary teeth. There is a minimum of 22 maxillary teeth and 27 dentary teeth in jaws of the largest well-preserved individuals known to date. Both dentitions (upper and lower) are bowed medially and are sinuous longitudinally. Maxillary and dentary crowns are tilted lingually on their roots, trapezoidal in outline and have crenellate (coarsely denticulate) margins. Adjacent crowns of teeth have mesiodistally (anteroposteriorly) expanded bases that overlap slightly and are consequently arranged en echelon. The dentitions are flanked by deep cheek pouches. Tooth abrasion is usually discontinuous along the dentition. In one individual nearly all teeth seem to be fully emerged and there is little evidence of abrasion. There is no physical evidence of a predentary, but the presence of this (typically ornithischian) element may be inferred from the structure of the symphyseal region of the dentary. The external narial and antorbital fenestrae are comparatively small, whereas the orbit and temporal fenestrae are large and open. A sclerotic ring was undoubtedly present and supported the eyeball, but it is too poorly preserved to allow it to be reconstructed with accuracy. A prominent supraorbital brow ridge overhangs the orbit. There are three osteoderms: palpebral, middle supraorbital and posterior supraorbital, sutured to the dorsal margin of the orbit. The occiput provides an area for attachment of a pair of curved, keratin-sheathed, osteodermal horns.Epistyloid bones project from the ventrolateral region of the braincase; their distal ends flank the anterolateral region of the neck. Rugose facets on either side of the basioccipital are suggested to have provided attachment sites for the epistyloid bones. Internally, the skull has a deeply vaulted snout and the nasal chambers are roofed by what are here named epivomer bones that appear to have been sutured to the dorsolateral edges of the vomers. Unusually, among dinosaurs generally, an epipterygoid is preserved attached to the dorsolateral surface of the pterygoid; there is no obvious point of articulation for the epipterygoid against the lateral wall of the braincase. A deep pit on the posterior surface of the quadrate of an immature specimen is suggestive of the existence of a remnant of cranial pneumatism. This pit becomes occluded in larger, more mature specimens.

Osteology of the archosauromorph Teyujagua paradoxa and the early evolution of the archosauriform skull

Abstract Archosauriformes are a major group of fossil and living reptiles that include the crown group Archosauria (birds, crocodilians and their extinct relatives) and closely related taxa. Archosauriformes are characterized by a highly diagnostic skull architecture, which is linked to the predatory habits of their early representatives, and the development of extensive cranial pneumaticity associated with the nasal capsule. The evolution of the archosauriform skull from the more plesiomorphic configuration present ancestrally in the broader clade Archosauromorpha was, until recently, elusive. This began to change with the discovery and description of Teyujagua paradoxa, an early archosauromorph from the Lower Triassic Sanga do Cabral Formation of Brazil. Here, we provide a detailed osteological description of the holotype and, thus far, only known specimen of T. paradoxa. In addition to providing new details of the anatomy of T. paradoxa, our study also reveals an early development of skull pneumaticity prior to the emergence of the antorbital fenestra. We use these new data to discuss the evolution of antorbital openings within Archosauriformes. Reappraisal of the phylogenetic position of T. paradoxa supports previous hypotheses of a close relationship with Archosauriformes. The data presented here provide new insights into character evolution during the origin of the archosauriform skull.

Psittacosaurus amitabha, a New Species of Ceratopsian Dinosaur from the Ondai Sayr Locality, Central Mongolia

Psittacosaurus is the most speciose nonavian dinosaur genus, represented by at least 10 (and possibly as many as 19) species uncovered over a wide geographic range. Here, we report a new species of large-bodied Psittacosaurus from the Ondai Sayr locality in central Mongolia, which has hitherto produced only one other Psittacosaurus specimen. This new species is characterized by: (1) an elongate snout, with a gently inclined rostronasal margin, (2) a cranium dorsally convex rather than flat, (3) a subtemporal length less than 40% of total skull length, (4) a maxillary lamina that cups the posterior toothrow, (5) five premaxillary foramina arrayed in an arc, (6) an antorbital fossa as long as it is wide, and (7) a palpebral with a well-developed posterior tonguelike process. Psittacosaurus amitabha is resolved as the most basal member of the genus Psittacosaurus in our phylogenetic analysis. This taxon expands our knowledge of the already-speciose genus Psittacosaurus as well as our understanding of the Ondai Sayr fauna, which is poorly known in comparison to other Mongolian Early Cretaceous localities.

New Proterochampsid Specimens Expand the Morphological Diversity of the Rhadinosuchines of the Chañares Formation (Lower Carnian, Northwestern Argentina)

The proterochampsids are crocodile-like, small- to medium-sized, quadrupedal and predatory stem-archosaurs. The rhadinosuchines are a clade of deeply nested proterochampsids that are the most abundant diapsids of the Massetognathus-Chanaresuchus Assemblage Zone of the Chanares Formation (lower Carnian) of north-western Argentina. Here, we describe two new rhadinosuchine proterochampsid specimens recently collected from the Massetognathus-Chanaresuchus Assemblage Zone of the Chanares Formation. These specimens are represented by an articulated partial skeleton and disarticulated cranial and postcranial bones, respectively. One of them is less complete, but its well-preserved disarticulated bones considerably expand the anatomical knowledge of certain regions of the rhadinosuchine skeleton. The combination of character-states present in both specimens and shared with other known rhadinosuchine species (e.g., Chanaresuchus bonapartei, Gualosuchus reigi, Rhadinosuchus gracilis) results in an unexpected increase of the morphological diversity of the group (e.g., antorbital fossa on the horizontal process of maxilla, well-exposed antorbital fossa on lacrimal, X-shaped osteoderms). We conclude that the taxonomy of the currently known rhadinosuchine specimens should be revisited, including a careful reappraisal of the intraspecific variation and the post-mortem deformation that affected the specimens in different ways.

A new Theriosuchus-like crocodyliform from the Maastrichtian of Romania

We describe a new, small-sized atoposaurid crocodyliform from the Upper Cretaceous (Maastrichtian) of Haţeg Basin, Romania. Aprosuchus ghirai gen. et sp. nov. is represented by a nearly complete three-dimensional skull with articulating mandibles and an associated fragmentary procoelous cervical vertebra that may have belonged to the same subadult individual. The new taxon possesses a heterodont dentition with at least four distinct tooth morphotypes including pseudocaniniform, pseudoziphodont lanceolate, ziphodont lanceolate and ‘low-crowned’ teeth. It differs from Sabresuchus sympiestodon, the other Theriosuchus-like taxon from the same stratigraphic unit, in having less enlarged fang-like teeth, a ‘w’ shaped naso-frontal articulation and extremely large palpebrals strongly fused to the orbital margin. The new taxon shares with other atoposaurids a broad oreinirostral rostrum, a vascular opening on the dorsal surface of postorbital bar, a small antorbital fenestra, a partially septated choanal groove and symmetrically developed lateral compression of maxillary teeth. Other synapomorphies of Aprosuchus are shared by a broader range of taxa including Wannchampsus and paralligatorids. The fossil record indicates that these small, highly specialized crocodylomorphs survived up to the latest Cretaceous by colonizing remote and rather complex insular ecosystems in the western Tethyan archipelago.

Ontogeny and variation of a protoceratopsid dinosaur Bagaceratops rozhdestvenskyi from the Late Cretaceous of the Gobi Desert

ABSTRACT Bagaceratops rozhdestvenskyi is a ceratopsian dinosaur from the Late Cretaceous Baruungoyot Formation of the Gobi Desert, closely related to Protoceratops spp. Several Bag. rozhdestvenskyi skulls demonstrate a wide range of variation in their morphology and size. Here I argue that the observed variability is most likely of intraspecific nature. Specimens classified in a few allegedly distinct species from the same or near-contemporary sediments, namely Gobiceratops minutus, Lamaceratops tereschenkoi and Platyceratops tatarinovi from Baruungoyot Formation, and Magnirostris dodsoni from Bayan Mandahu, are younger subjective synonyms of Bag. rozhdestvenskyi. They plausibly represent an ontogenetic series within the latter. Breviceratops kozlowskiiis a distinct taxon. The evolutionary relationships within Protoceratopsidae are complicated by the mosaic distribution of plesiomorphic and derived features in distinct species. I suggest that taxa distribution and observed changes in morphology are an evidence for the ancestral position of Protoceratops andrewsi among protoceratopsids. It implies possible temporary separation between the geological formations of the Gobi Desert yielding distinct protoceratopsid species. The novel evolutionary scenario suggests number of convergences that occurred in Protoceratopsidae and Ceratopsoidea (reduction of the premaxillary dentition, fusion of nasals, development of the accessory antorbital fenestra). Present study reveals the significance of the intraspecific and ontogenetic variation in the study of the neoceratopsian taxonomy.