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Abstract

Skorpiovenator bustingorryi is a derived abelisaurid theropod represented by a fairly complete skeleton from the Late Cretaceous sedimentary beds of north-western Patagonia. Although some features were described in the original paper, mainly related to the skull, the appendicular anatomy remains undescribed. The aim of the present contribution is to provide a detailed description and analysis of the available appendicular bones, including comparisons with other ceratosaurian theropods close to Skorpiovenator. In this way, new autapomorphies emerged to further distinguish Skorpiovenator from its relatives. Furthermore, a comprehensive phylogenetic analysis was performed and several characteristics of the hind limb, in particular some of the autopodium, resulted in the identification of new apomorphic traits for Ceratosauria and Abelisauridae. These features might prove to be useful for future phylogenetic analyses and may help to resolve the still confusing and debated internal relationships of abelisaurid theropods.

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Abelisaurids are among the most abundant and diverse Patagonian Late Cretaceous theropods. Here, we present a new furileusaurian abelisaurid, Llukalkan aliocranianus gen. et sp. nov., represented by cranial remains from the Bajo de la Carpa Formation (Santonian) at La Invernada fossil area, northwestern Patagonia. Features characterizing this taxon include a possible caudal tympanic recess posterior to the columellar recess, a T-shaped lacrimal with jugal ramus lacking a suborbital process, and large foramina for caudal middle cerebral veins widely separated from the median supraoccipital crest. In addition to this, a bulge on the anteromedial border of the supratemporal fossa, tall and posteriorly projected paroccipital processes, basal tubera interconnected distally, a triangular basisphenoid recess, and a single foramen for the sphenoidal artery on the basisphenoid, differentiate Llukalkan from Viavenator exxoni. The latter is the other furileusaurian taxon from the same area and stratigraphic unit. Although the holotype of Llukalkan probably corresponds to a sub-adult—as the lacrimal morphology suggests— the possibility that it represents a juvenile of V. exxoni is discarded based mainly on the presence of a caudal tympanic recess (which is absent in V. exxoni). The probable coexistence of two abelisaurid taxa demonstrates that the abelisaurids were one of the most important—and likely the main—predator component of the ecosystems, not only in this area, but also in all of Patagonia, during the Late Cretaceous.
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Xenotarsosaurus bonapartei was the third abelisaurid theropod dinosaur to be named from Argentina. The holotype comprises two partial anterior dorsal vertebrae and a complete right hind limb from the Upper Cretaceous (lower Cenomanian–upper Turonian) Bajo Barreal Formation, central Patagonia, Argentina. The materials display morphological features that undoubtedly position Xenotarsosaurus within Abelisauroidea. Moreover, detailed comparisons with members of that theropod group confirm the close relationship of this taxon to abelisaurids. Here we provide an emended diagnosis of Xenotarsosaurus bonapartei that includes five newly recognized autapomorphies: (1) anterior dorsal vertebrae with large, strongly dorsoventrally developed parapophyses; (2) anterior dorsal vertebrae with well-developed centroprezygapophyseal fossae that are taller dorsoventrally than wide mediolaterally; (3) fibular condyle of femur triangular in shape and projecting posteriorly; (4) well-marked groove on the anterolateral corner of the proximal fibula; and (5) iliofibularis tubercle of fibula distally interrupted by a hook-like shaped concavity. To determine its systematic position within Abelisauroidea, we incorporated Xenotarsosaurus into a phylogenetic analysis, recovering this theropod as a non-carnotaurine abelisaurid more derived than Eoabelisaurus mefi. Xenotarsosaurus displays several plesiomorphic traits when compared with penecontemporaneous abelisaurids from the Neuquén Group. Similarly, other non-avian dinosaur taxa from the Bajo Barreal Formation are frequently postulated as more phylogenetically basal than coeval forms from northern Patagonia. This scenario suggests the potential existence of provincialism in early Late Cretaceous continental vertebrate faunas of southern South America. The present study increases knowledge of abelisaurid systematics, evolution, and paleobiogeography and augments our understanding of the Late Cretaceous dinosaur assemblage of central Patagonia.
Article
Abelisaurid theropods are well known in Cretaceous beds along South America, particularly Patagonia. However, the record of latest Cretaceous abelisauroids is still far from satisfactory. Until recently, few taxa were described from Maastrichtian beds: Carnotaurus sastrei and Quilmesaurus curriei, but also perhaps Abelisaurus comahuensis. In this contribution, we describe a new genus and species (Niebla antiqua gen. et sp. nov.), of medium-sized abelisaurid coming from Late Cretaceous (Maastrichtian) beds from Río Negro province, northern Patagonia, Argentina. The paleohistological analysis shows that this individual has reached the somatical maturity and that it represents an adult of a mid-sized abelisaurid. The specimen is represented by a nearly complete braincase, fragmentary jaw and teeth, relatively complete scapulocoracoid, dorsal ribs and incomplete vertebrae. The new taxon is relatively small, much smaller than other coeval abelisaurids such as Carnotaurus and Abelisaurus. The braincase shows autapomorphic features such a dorsoventrally tall basal tuber and postemporal foramen enclosed by parietal and exoccipitals. The scapulocoracoid is notably similar to that of Carnotaurus in having a posterodorsally oriented glenoid, a dorsoventrally expanded and wide coraco-scapular plate, and a very narrow and straight scapular blade. These features are very different from those of other abelisaurids, which may indicate a unique conformation of the pectoral girdle among these South American theropods.
Article
Abelisaurids were one of the most successful theropod dinosaurs during Cretaceous times. They are featured by numerous derived skull traits, such as heavily ornamented bones, short and tall snout, and a strongly thickened cranial roof. Furthermore, nasals are distinctive on having two distinct nasal patterns: strongly transversely convex and heavily sculptured (e.g., Carnotaurus), and transversely concave, with marked bilateral crests and poorly sculptured surfaces (e.g., Rugops). Independently of the pattern , some abelisaurid nasals (e.g., Rugops) show a distinctive row of large foramina on the dorsal surface, which were in general associated to skin structures (scales). Skorpiovenator bustingorryi is a derived abelisaurid coming from the upper Cretaceous beds of northwestern Patagonia, represented by an almost complete skeleton including a well-preserved skull. Particularly, the skull of Skorpiovenator shows nasal bones characterized by being transversely concave, rimmed by lateral crests and with a conspicuous row of foramina on the dorsal surface. But more interesting is that the skull roof also exhibits a row of large foramina that seem to be continuous with the previous nasal foramina. CT scans made on the skull corroborates a novel feature within theropods: the nasal foramina on the external surface are linked to an internal canal that runs across the nasal bones. We compared this feature with CT scans of Carnotaurus and revealed that it also possess an internal system as in Skorpiovenator, but being notably smaller. The symmetry and disposition of the foramina in the nasal and skull roof bones of Skorpiovenator would indicate a neurovascular correlate (i.e., blood vessels and nerves), probably to the lateral nasal and supraorbital vessels and the trigeminal nerve. The biological significance of such neurovascular system can be conjectured from several hypotheses. A possible one involves an enhanced blood volume in these bones linked to a zone of thermal exchange, which may help avoid overheat of encephalic tissues. Another plausible hypothesis takes into account the presence of display skin structures in which blood volume nourished the mineralized skin, which would have a role in intraspecific communication. However, other more speculative explanations should not be discarded such as a correlation with integu-mentary sensory organs.
Article
A new ceratosaurian theropod dinosaur, Huinculsaurus montesi gen. et sp. nov., is described here. This taxon is based on the last three dorsal vertebrae and the first and second sacral vertebrae found in association at Aguada Grande, Neuquén Province, Argentina. Although fragmentary, Huinculsaurus shows a unique mix of features which differentiates it from all other theropods, including the sympatric abelisaurid Ilokelesia, and is diagnosed by prezygapophyseal articular facets twice longer than wide, anterior centroparapophyseal lamina strongly developed as an extensive lateral lamina in the posterior dorsal vertebrae, pneumatic foramina located ventrally to the postzygodiapophyseal lamina in the posterior dorsal vertebrae, posteriorly tapering postzygapophysis pointed posteriorly, and an accessory lamina bisecting the parapophyseal centrodiapophyseal fossa in the posterior dorsal neural arches. Phylogenetic analyses recovered Huinculsaurus as most closely related to the Late Jurassic Elaphrosaurus than to other Cretaceous abelisauroids, suggesting the persistence of the elaphrosaurine lineage in South America up to the early Late Cretaceous.
Article
Phylogenetic relationships of megaraptorid theropods are under intense debate. Some authors interpret them as archaic allosauroids that survived up to Late Cretaceous, whereas others consider megaraptorids as basal tyrannosauroids. The recently described Patagonian taxon Murusraptor barrosaensis offers novel information on skull, axial and hind limb anatomy, all of which may help in elucidating the phylogenetic affinities of megaraptorids as a whole. Murusraptor is particularly similar to juvenile specimens of tyrannosaurids; both share: 1) lacrimal with a long anterior process; 2) corneal process and; 3) lateral pneumatic fenestra; 4) square and dorsoventrally low frontals; 5) parietals with well-developed sagittal and nuchal crests, among other features. The current study lends further support to the hypothesis that megaraptorans are basal members of Coelurosauria (supported by 20 synapomophies), with strongest affiliation with Tyrannosauroidea (supported by >20 synapomorphies).
Article
Several remains of an abelisaurid theropod including a nearly complete sacral complex articulated with both ilia, the distal boot of the pubes, the furcula, teeth, and fragments of transverse processes of caudal vertebrae was discovered in the La Invernada fossil site, northern Patagonia from the Bajo de la Carpa Formation (Santonian). The sacrum exhibit features typical of abelisauroids as narrowing of the middle sacral centra, fused neural spines forming a continuous sheet and ventral bow of the sacrum, in lateral view. The furcula represents de first mention of this bone for a South American abelisaurid. The morphology observed in the transverse processes with distal end projected forward and sagittal ridge on the ventral surface allows its taxonomic assignment to the clade Brachyrostra.
Article
Abelisaurid theropods were most abundant in the Gondwana during the Cretaceous Period. Pycnonemosaurus nevesi was the first abelisaurid dinosaur described from the Bauru Group (Brazil, Upper Cretaceous). Nevertheless, its initial description was based on the comparison of a restricted number of remains with other abelisaurids. In this paper, I present a new description of the morphology of Pycnonemosaurus nevesi, including three new caudal transverse processes and a discussion of several new characteristics based on perspectives derived from recently described abelisauroids. Pycnonemosaurus nevesi differs from other abelisaurids based on the following features: a pubis with a small rounded foot and a ventrally-bowed anterior distal end; posterior caudal vertebrae with a hook-shaped transverse process that has an anterodistal expansion that is short and bowed; a strong and massive tibia with a well-developed lateral malleolus that is ventrally expanded. The unfused sutures represent signs of skeletal immaturity, but the specific ontogenetic stage is still uncertain. The current phylogenetic analysis suggests strongly relationship within Pycnonemosaurus and the most-derived abelisaurids (e.g Carnotaurus and Aucasaurus).
Article
A partial postcranial skeleton from the Late Jurassic (Tithonian) of Dorset, England represents a new species of the theropod dinosaur Stokesosaurus, Stokesosaurus langhami. S. langhami is a member of Tyrannosauroidea, showing a distinct median vertical ridge on the lateral surface of the ilium, a prominent shelf medial to the preacetabular notch, a pronounced ischial tubercle, and a tibia that is elongate relative to the femur. One of only two definitive Jurassic tyrannosauroids known from more than isolated elements, it is the largest Jurassic tyrannosauroid reported to date and provides additional evidence for the presence of relatively small-or medium-sized basal tyrannosauroids in Asia, North America, and Europe during the Late Jurassic. The occurrence of Stokesosaurus in the Tithonian of the UK and USA and the absence of tyrannosauroids in contemporaneous west African faunas supports the hypothesis of a paleobiogeographic link during the Late Jurassic between North America and Europe, to the exclusion of Africa.
Article
Ontogenetic variation is documented within many dinosaur species, but extreme ontogenetic changes are rare among dinosaurs, particularly among theropods. Here, we analyze 19 specimens of the Jurassic ceratosaurian theropod Limusaurus inextricabilis, representing six ontogenetic stages based on body size and histological data. Among 78 ontogenetic changes we identify in these specimens, the most unexpected one is the change from fully toothed jaws in the hatchling and juvenile individuals to a completely toothless beaked jaw in the more mature individuals, representing the first fossil record of ontogenetic edentulism among the jawed vertebrates. Jaw morphological data, including those derived from Mi-CT and SR-μCT scanning of Limusaurus specimens, reveal dental alveolar vestiges and indicate that ontogenetic tooth loss in Limusaurus is a gradual, complex process. Our discovery has significant implications for understanding the evolution of the beak, an important feeding structure present in several tetrapod clades, including modern birds. This radical morphological change suggests a dietary shift, probably from omnivory for juvenile Limusaurus to herbivory for adult Limusaurus, which is also supported by additional evidence from gastroliths and stable isotopes. Incorporating new ontogenetic information from Limusaurus into phylogenetic analyses demonstrates surprisingly little effect on its placement when data from different stages are used exclusively, in contrast to previous analyses of tyrannosaurids, but produces subtle differences extending beyond the placement of Limusaurus.
Article
Here, we describe new theropod materials (several isolated teeth, an axis, two caudal centra and a proximal left tibia) from the type locality of the Bajada Colorada Formation (Lower Cretaceous, Berriasian–Valanginian). Although fragmentary, the recovered material shows a diverse association of meat-eating dinosaurs for this poorly understood period of time. Three techniques were used to assess the phylogenetic position of the isolated teeth: multivariate (PCA), discriminant and phylogenetic analyses. The morphotypes 1, 2 and 3 (small non-recurved isolated crowns) were regarded as Theropoda indet., as our analyses failed to support a more precise classification. Two large almost complete ziphodont crowns, considered morphotype 4, were identified as belonging to megalosaurid tetanurans by phylogenetic, discriminant and multivariate analyses, thus likely representing the first record of this ancient family in South America, and the youngest worldwide. We refer the axis to a small abelisauroid ceratosaurian based on the following suite of characters; long and pointed epipophyses, a pneumatic foramen in the centrum, the invaginated spinopostzygapophyseal lamina, and the anteroposteriorly long, dorsally convex, and dorsally/posteriorly unexpanded neural spine. The caudal centra can be referred to a theropod, and tentatively to Abelisauroidea, suggesting that they may belong to the same taxon than that represented by the axis. The tibia lacks the incisura tibialis, presents a low, sharp and proximally positioned fibular crest, and has a deep lateral fossa, which is limited dorsally by a coarse, rounded in section and anteriorly directed crest. This combination of features allows to refer it to a large abelisaurid. The Bajada Colorada dinosaur record includes so far a small abelisauroid, a large abelisaurid, a probably medium to large megalosaurid tetanuran, diplodocid and dicraeosaurid sauropods. It shows some similarities with Middle and Upper Jurassic units in central Patagonia, Africa and Portugal, suggesting that no significant dinosaur faunal turnover took place through the Jurassic-Cretaceous event in the southern part of South America.
Article
Version 1.5 of the computer program TNT completely integrates landmark data into phylogenetic analysis. Landmark data consist of coordinates (in two or three dimensions) for the terminal taxa; TNT reconstructs shapes for the internal nodes such that the difference between ancestor and descendant shapes for all tree branches sums up to a minimum; this sum is used as tree score. Landmark data can be analysed alone or in combination with standard characters; all the applicable commands and options in TNT can be used transparently after reading a landmark data set. The program continues implementing all the types of analyses in former versions, including discrete and continuous characters (which can now be read at any scale, and automatically rescaled by TNT). Using algorithms described in this paper, searches for landmark data can be made tens to hundreds of times faster than it was possible before (from T to 3T times faster, where T is the number of taxa), thus making phylogenetic analysis of landmarks feasible even on standard personal computers.
Article
Theropod dinosaurs from the Late Jurassic of Gondwana are still poorly known, with Elaphrosaurus bambergi Janensch, 1920, from the late Kimmeridgian of Tendaguru, Tanzania, being the only taxon represented by more than isolated remains from Africa. Having long been considered a coelurosaurian, more specifically an ornithomimosaur, Elaphrosaurus is currently regarded as a basal ceratosaur. Here, we revise the osteology and phylogenetic position of this important taxon. Elaphrosaurus shows many unusual osteological characters, including extremely elongated and constricted cervical vertebrae, an expansive shoulder girdle with strongly modified forelimbs, a relatively small ilium, and elongate hindlimbs with a very small ascending process of the astragalus that is fused to the tibia. We found this taxon to share many derived characters with noasaurids, such as: strongly elongate cervical and dorsal vertebrae; low, rectangular neural spines in the mid-caudal vertebrae; presence of only an anterior centrodiapophyseal lamina in anterior caudal vertebrae; presence of a wide, U-shaped notch between the glenoid and the anteroventral hook in the coracoid; a laterally flared postacetabular blade of the ilium; a flat anterior side of the distal tibia; and a reduced shaft of metatarsal II. Our analysis placed Elaphrosaurus within a dichotomous Noasauridae as part of a Jurassic subclade, here termed Elaphrosaurinae, that otherwise includes taxa from eastern Asia. These results underscore the long and complex evolutionary history of abelisauroids, which is still only beginning to be understood.
Article
The emergence of a reversed hallux (first pedal digit) in the theropod lineage is regarded as an important indicator of increasing grasping ability and possibly arboreality. However, functions of the pes with a plesiomorphic, non-reversed hallux in non-avian theropods, other than the major role in supporting and propelling the body during the stance phase, have not been fully addressed. To clarify hallucal function, 21 specimens of non-avian theropods were examined in this study. Ancestrally in Theropoda, the proximal end of the first metatarsal reached the ankle joint as in extant crocodilians. The size of the first metatarsal was then reduced, lost direct contact with the ankle joint, and shifted more plantarly in basal neotheropods. In articulated coelophysoid specimens, the proximal end of the first metatarsal attaches to the medioplantar margin of the second metatarsal, whereas the distal part of the first metatarsal and phalanges of the hallux lies on the plantar aspect of the pes. In four articulated specimens of the dromaeosaurid Velociraptor, preserved positions of the first metatarsal on the second metatarsal vary from the medial to plantar aspects, and correspondingly the rotational axis of the extension/flexion movement at its distal articulation ranges from mediolateral to dorsoplantar. Similar conditions are also observed in troodontid specimens, suggesting the presence of intermetatarsal mobility between the first and second metatarsals enabled by a convex proximal articular surface of the first metatarsal in deinonychosaurians similar to the one present in extant birds. Citation for this article: Hattori, S. 2016. Evolution of the hallux in non-avian theropod dinosaurs. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1116995.
Article
Tyrannosauroids are among the most distinctive and best known of Mesozoic theropods. Diagnostic skeletal material for Tyrannosauridae is at present limited to the last part of the Late Cretaceous of eastern and central Asia (China and Mongolia) and North America; the more inclusive Tyrannosauroidea includes taxa from the Late Jurassic of Europe and North America and the Early Cretaceous of Europe and Asia. Tyrannosaurids include some of the largest known theropod taxa (up to 13 m long and weighing six or more tonnes). The five most completely known species (Tyrannosaurus rex, Tarbosaurus bataar, Daspletosaurus torosus, Albertosaurus sarcophagus, and Gorgosaurus libratus) are all represented by individuals with femora more than 100 cm in length, reaching 138 cm in the largest Tyrannosaurus rex.
Article
The Neuquén Group conform a succession of non-marine deposits of fluvial, aeolian and shallow lacustrine origin, developed in the Neuquén Basin between the Lower Cenomanian and the Middle Campanian (Upper Cretaceous). This succession reaches 1,200 m of maximum thick, being bounded at the base and the top by the Patagonidican and Huantraiquican unconformities respectively. In this paper two new formations are proposed (Los Bastos and Sierra Barrosa Formations, nom. nov.), and the subgroups were reconfigurated. Consequently, the Neuquén Group is composed lithostratigraphically by three subgroups and nine formations. The Río Limay Subgroup comprehend the Candeleros and Huincul Formations, representing low sinuosity channel deposits developed under tectonic and climatic subordinated control. The Río Neuquén Subgroup comprehend to the Cerro Lisandro, Portezuelo, Los Bastos, Sierra Barrosa and Plottier Formations, conforming a psamitic and pelitic interstratified succession of fluvial origin, mainly developed under climatic control. Finally, the Río Colorado Subgroup integrated by the Bajo de la Carpa and Anacleto Formations, comprehend fluvial deposits developed under a new dominant tectonic control. At the start of the deposition of the Neuquén Group, the Dorsal de Huincul worked as a structural high, generating toward the south of the Neuquén Basin one a small endorreic depocentre corresponding to the Picún Leufú Sub-basin, scheme that would have been keep up to ends of the deposit of the Candeleros Formation. During the rest of the deposit of this sequence, the drainage would have been kept without major modifications towards the pacific slope across a central collector flowing in sense N-NW. Towards the end of the deposit of the Neuquén Group takes place the inversion of the regional slope of the basin, making possible the ingression of the Atlantic Ocean and the beginning of the deposition of the Malargüe Group.