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Appendicular osteology of Skorpiovenator bustingorryi (Theropoda, Abelisauridae) with comments on phylogenetic features of abelisaurids
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.
... In other ceratosaurs, the pelvic bones are at least partially fused around the acetabulum (e.g. Pol and Rauhut, 2012;Rauhut and Carrano, 2016;Cerroni et al., 2022). ...
... As in several previously published phylogenies Cerroni et al., 2022), there are several terminal taxa that form the basal branches of Abelisauridae (Eoabelisaurus, Rugops, Spectrovenator) and two major clades: Brachyrostra and 'Majungasaurinae' (Fig. 18). Brachyrostra includes derived Late Cretaceous forms from South America and possibly Europe, whereas 'Majungasaurinae' includes abelisaurids from India and Madagascar. ...
... Another difference from the parsimony results is that Xenotarsosaurus is allied with other Cretaceous taxa rather than with Eoabelisaurus (Fig. 19a). These relationships contradict both the results obtained from the time-free methods (parsimony and nonclock Bayesian inference) as well as from all previous iterations of this dataset (Carrano and Sampson, 2008;Pol and Rauhut, 2012;Rauhut and Carrano, 2016;Baiano et al., 2021Ibiricu et al., 2021;Cerroni et al., 2022). However, the unorthodox placements of the taxa mentioned above are very weakly supported in the tip-dating analysis (with PP ≤ 0.26; Fig. 19a) and there is a considerable topological uncertainty across the whole tree (average PP = 0.237; Fig. 20). ...
Gondwanan dinosaur faunae during the 20 Myr preceding the Cretaceous-Palaeogene (K/Pg) extinction included several line-ages that were absent or poorly represented in Laurasian landmasses. Among these, the South American fossil record contains diverse abelisaurids, arguably the most successful groups of carnivorous dinosaurs from Gondwana in the Cretaceous, reaching their highest diversity towards the end of this period. Here we describe Koleken inakayali gen. et sp. n., a new abelisaurid from the La Colonia Formation (Maastrichtian, Upper Cretaceous) of Patagonia. Koleken inakayali is known from several skull bones, an almost complete dorsal series, complete sacrum, several caudal vertebrae, pelvic girdle and almost complete hind limbs. The new abelisaurid shows a unique set of features in the skull and several anatomical differences from Carnotaurus sas-trei (the only other abelisaurid known from the La Colonia Formation). Koleken inakayali is retrieved as a brachyrostran abeli-saurid, clustered with other South American abelisaurids from the latest Cretaceous (Campanian-Maastrichtian), such as Aucasaurus, Niebla and Carnotaurus. Leveraging our phylogeny estimates, we explore rates of morphological evolution across ceratosaurian lineages, finding them to be particularly high for elaphrosaurine noasaurids and around the base of Abelisauridae, before the Early Cretaceous radiation of the latter clade. The Noasauridae and their sister clade show contrasting patterns of morphological evolution, with noasaurids undergoing an early phase of accelerated evolution of the axial and hind limb skeleton in the Jurassic, and the abelisaurids exhibiting sustained high rates of cranial evolution during the Early Cretaceous. These results provide much needed context for the evolutionary dynamics of ceratosaurian theropods, contributing to broader understanding of macroevolutionary patterns across dinosaurs.
... To test the phylogenetic position of Aucasaurus based on new axial information, we carried out an analysis based on the most recently studies of Ceratosauria (Tortosa et al., 2014;Filippi et al., 2016;Rauhut & Carrano, 2016;Baiano, Coria & Cau, 2020;Baiano et al., 2021Aranciaga Rolando et al., 2021;Cerroni et al., 2022). We added 11 characters (seven new and four from other sources) to the data matrices of and Cerroni et al. (2022); we also added three new taxa (i.e., Kurupi,Thanos,and MPM 99). ...
... To test the phylogenetic position of Aucasaurus based on new axial information, we carried out an analysis based on the most recently studies of Ceratosauria (Tortosa et al., 2014;Filippi et al., 2016;Rauhut & Carrano, 2016;Baiano, Coria & Cau, 2020;Baiano et al., 2021Aranciaga Rolando et al., 2021;Cerroni et al., 2022). We added 11 characters (seven new and four from other sources) to the data matrices of and Cerroni et al. (2022); we also added three new taxa (i.e., Kurupi,Thanos,and MPM 99). The resulting data matrix consisted of 246 characters and 46 taxa (Data S1). ...
Aucasaurus garridoi is an abelisaurid theropod from the Anacleto Formation (lower Campanian, Upper Cretaceous) of Patagonia, Argentina. The holotype of Aucasaurus garridoi includes cranial material, axial elements, and almost complete fore- and hind limbs. Here we present a detailed description of the axial skeleton of this taxon, along with some paleobiological and phylogenetic inferences. The presacral elements are somewhat fragmentary, although these show features shared with other abelisaurids. The caudal series, to date the most complete among brachyrostran abelisaurids, shows several autapomorphic features including the
presence of pneumatic recesses on the dorsal surface of the anterior caudal neural arches, a tubercle lateral to the prezygapophysis of mid caudal vertebrae, a marked protuberance on the lateral rim of the transverse process of the caudal vertebrae, and the presence of a small ligamentous scar near the anterior edge of the dorsal surface in the anteriormost caudal transverse process. The detailed study of the axial skeleton of Aucasaurus garridoi has also allowed us to identify characters that could be useful for future studies attempting to resolve the internal phylogenetic relationships of Abelisauridae. Computed tomography scans of some caudal vertebrae show pneumatic traits in neural arches and centra, and thus the first reported case for an abelisaurid taxon. Moreover, some osteological correlates of soft tissues present in Aucasaurus and other abelisaurids, especially derived brachyrostrans, underscore a previously proposed increase in axial rigidity within Abelisauridae.
... In lateral view, the cnemial crest is projected anterodorsally so that it rises well above the condyles, as is typical of Ceratosauria (Ju arez- Valieri et al., 2007;Carrano and Sampson, 2008). The dorsal projection seen here is comparable to that of Genusaurus (Accarie et al., 1995), Quilmesaurus (Coria, 2001), Aucasaurus (Coria et al., 2002), and Skorpiovenator (Cerroni et al., 2022). The dorsal projection is less well-developed than in Indosuchus (Novas et al., 2004), Majungasaurus (Carrano, 2007), Pycnocnemisaurus (Kellner and Campos, 2002) and an abelisaurid from the Marilia Formation of Brazil (Machado et al., 2013). ...
... The lateral condyle is broad and rounded. The medial condyle is narrower mediolaterally than in Skorpiovenator (Cerroni et al., 2022), Xenotarsosaurus (Martínez et al., 1986), or an unnamed abelisaurid from the Late Cretaceous Marilia Formation of Brazil (Machado et al., 2013). This feature may be exaggerated by crushing of the proximal end of the bone, but Majungasaurus (Carrano, 2007) has a similarly narrow tibia. ...
The end of the Cretaceous saw the evolution of endemic dinosaur faunas on different landmasses, driven by continental fragmentation. Understanding the evolution of these biogeographic patterns is important for understanding the evolution of Mesozoic ecosystems. However, the faunas of the southern land masses remain understudied relative to the intensively sampled dinosaur faunas of western North America and Asia. In particular, the latest Cretaceous of Africa remains largely unknown, with only a handful of taxa reported so far, including titanosaurian sauropods, the lambeosaurine Ajnabia odysseus, and the large abelisaurid theropod Chenanisaurus barbaricus. We report two new abelisaurid fossils from the upper Maastrichtian phosphates of the Ouled Abdoun Basin, in northern Morocco. The first is the tibia of a medium-sized abelisaurid from Sidi Chennane, with an estimated length of ~5 m. The tibia has a strongly hooked cnemial crest resembling that of the South American Quilmesaurus and Aucasaurus. The highly rugose bone texture suggest the animal was mature, rather than a juvenile of the larger Chenanisaurus. The second is a small right second metatarsal from Sidi Daoui,. The metatarsal measures 190 mm in length, suggesting a small animal, ~2.6 m in length. The metatarsal shows strong mediolateral compression, a feature present in noasaurids and some early abelisaurids, but absent in most Late Cretaceous abelisaurids. It is distinct from other abelisauroids in the strong constriction and bowing of the shaft in lateral view, and the medial curvature of the bone in anterior view. Bone texture suggests it comes from a mature individual. The small size, gracile proportions and unusual shape of the metatarsal suggest it is not closely related to other latest Cretaceous abelisaurids. The new fossils suggest as many as three abelisaurid taxa coexisted in the late Maastrichtian of Morocco, showing dinosaurs were highly diverse in North Africa prior to the end-Cretaceous mass extinction.
... This character is found in derived abelisaurids such as the furileusaurian Brachyrostra Carnotaurus sastrei [48], Aucasaurus garridoi [34] and Koleken inakayali [3]. In the less derived Majungasaurus crenatissimus [42] and Skorpiovenator bustingorryi [49], the dorsal margin of the ilium is convex, while it is possibly undulated in Rahiolisaurus gujaratensis [50] and Ekrixinatosaurus novasi [51]. Interestingly, a straight dorsal margin is also present in Genusaurus sisteronis, from the Albian of France, a possible furileusaurian (see below and [34]). ...
An articulated group of skeletal elements comprising a sacrum, both ilia and a first caudal vertebra, plus an isolated tooth found in immediate proximity to the bones, from the lower Cenomanian Chalk at Saint-Jouin-Bruneval (Seine-Maritime, Normandy, France) is described and attributed to a new genus and species of abelisaurid theropod, Caletodraco cottardi, on the basis of several characters of the sacrum and pelvis. The peculiar shape of the transverse process of the first caudal vertebra shows that Caletodraco cottardi differs from majungasaurine abelisaurids previously described from Europe, such as Arcovenator escotae, and belongs to the Furileusauria, a group of derived abelisaurids hitherto recognized only from South America. The presence of a furileusaurian abelisaurid in the Cenomanian of Normandy suggests that the biogeographical history of the Abelisauridae in Europe was more complex than hitherto admitted. Several previously described European abelisaurids, such as the Albian Genusaurus sisteronis, may in fact belong to the Furileusauria.
We present the pelvic and hindlimb musculature of the abelisaurid Skorpiovenator bustingorryi, constituting the most comprehensive muscle reconstruction to date in ceratosaur theropods. Using extant phylogenetic bracket method, we reconstructed 39 muscles that can commonly found in extant archosaurs. Through the identification of bone correlates, we recognized thigh and hindlimb muscles including knee extensors, m. iliofibularis, m. flexor tibialis externus, mm. caudofemorales, mm. puboischiofemorales, and crus muscles important in foot extension and flexion (e.g., m. tibialis anterior, mm. gastrocnemii). Also, autopodial intrinsic muscles were reconstructed whose function involve extension (m. extensor digiti 2–4), flexion (mm. flexor digitorum brevis superficialis), interdigital adduction (m. interosseus dorsalis) and abduction (m. interosseous plantaris, m. abductor 4). Abelisaurids like Skorpiovenator show a deep pre‐ and postacetabular blade of the ilia and enlarged cnemial crests, which would have helped increasing the moment arm of muscles related to hip flexion and hindlimb extension. Also, pedal muscles related to pronation were probably present but reduced (e.g., m. pronator profundus). Despite some gross differences in the autopodial morphology in extant outgroups (e.g., crocodilian metatarsus and avian tarsometatarsus), the present study allows us to hypothesize several pedal muscles in Skorpiovenator. These muscles would not be arranged in tendinous bundles as in Neornithes, but rather the condition would be similar to that of crocodilians with several layers formed by fleshy bellies on the plantar and dorsal aspects of the metatarsus. The musculature of Skorpiovenator is key for future studies concerning abelisaurid biomechanics, including the integration of functional morphology and ichnological data.
A small-bodied ornithomimosaur, Kinnareemimus khonkaenensis Buffetaut, Suteethorn, and Tong 2009, from the Lower Cretaceous Sao Khua Formation of northeastern Thailand was first reported in 1995, then named in 2009. It is the only report of this group in Southeast Asia and was concluded to be a member of Ornithomimosauria, more derived than Harpymimus and Garudimimus but more basal than Archaeornithomimus. Since then, a few published studies have analyzed the phylogenetic position of Kinnareemimus but failed to find it within the clade Ornithomimosauria. Here, the phylogenetic analyses for assessing the relationships of Kinnareemimus within Ornithomimosauria were performed. The results suggested it might be a basal ornithomimosaur or belongs to the subclade Deinocheiridae. Kinnareemimus shares with deinocheirids the proximally projected cnemial crest and the shape of the fibula in proximal view. Its metatarsal III shows subarctometatarsalian condition with metatarsal III pinched between II and IV but visible in anterior view proximally, which might have evolved independently from other ornithomimids. However, the basal position of Kinnareemimus could also be due to the immaturity and the incomplete nature of this animal.
Soft tissue reconstructions of fossil vertebrates provide valuable data and support for functional interpretations. Here, we present the pelvic and hindlimb muscular reconstruction of Buitreraptor gonzalezorum, a unenlagiine dromaeosaurid theropod. Dromaeosaurids employed their hindlimbs for locomotion and predation, with a specialised pedal digit II. Using the ‘extant phylogenetic bracket’ methodology, we reconstruct 29 muscles out of the 37 typically found in extant archosaur hindlimbs. Some characteristics resemble those of specialised cursorial tetrapods. For example, M. iliotibialis shows an increased hip abduction moment arm, while primary protractors exhibit an increased hip flexion moment arm, and main retractors display a decreased hip extension moment arm. However, positive allometric growth of
limb inertia relative to body mass and the gradual transition to a ‘knee-based’ locomotion observed in extant birds may have influenced muscle moment arms. Regarding predatory functions, the muscular features and length proportions of the metatarsus and pedal digit II phalanges suggest faster movements compared to derived dromaeosaurids. Also, a robust flexor muscle, likely M. flexor perforans et perforatus digiti II, was inserted on the proximoventral heel of the second phalanx of pedal digit II. These functional interpretations suggest the presence of specialised behaviours different from those observed in more derived dromaeosaurids.
Tyrannosaurs are among the most intensively studied and best-known dinosaurs. Despite this, their relationships and systematics are highly controversial. An ongoing debate concerns the validity of Nanotyrannus lancensis, interpreted either as a distinct genus of small-bodied tyrannosaur or a juvenile of Tyrannosaurus rex. We examine multiple lines of evidence and show that the evidence strongly supports recognition of Nanotyrannus as a distinct species for the following reasons: 1. High diversity of tyrannosaurs and predatory dinosaurs supports the idea that multiple tyrannosaurids inhabited the late Maastrichtian of Laramidia; 2. Nanotyrannus lacks characters supporting referral to Tyrannosaurus or Tyrannosaurinae but differs from T. rex in >150 morphological characters, while intermediate forms combining the features of Nanotyrannus and T. rex are unknown; 3. Histology shows specimens of Nanotyrannus showing (i) skeletal fusions, (ii) mature skull bone textures, (iii) slow growth rates relative to T. rex, (iv) decelerating growth in their final years of life, and (v) growth curves predicting adult masses of~1500 kg or less, showing these animals are subadults and young adults, not juvenile Tyrannosaurus; 4. growth series of other tyrannosaurids, including Tarbosaurus and Gorgosaurus, do not show morphological changes proposed for a Nanotyrannus-Tyrannosaurus growth series, and deriving Tyrannosaurus from Nanotyrannus requires several changes inconsistent with known patterns of dinosaur development; 5. Juvenile T. rex exist, showing diagnostic features of Tyrannosaurus; 6. Phylogenetic analysis suggests that Nanotyrannus may lie outside Tyrannosauridae. Tyrannosaur diversity before the K-Pg extinction is higher than previously appreciated. The challenges inherent in diagnosing species based on fossils mean paleontologists may be systematically underestimating the diversity of ancient ecosystems.
Piatnitzkysauridae were Jurassic theropods that represented the earliest diverging branch of Megalosauroidea, being one of the earliest lineages to have evolved moderate body size. This clade's typical body size and some unusual anatomical features raise questions about locomotor function and specializations to aid in body support; and other palaeobiological issues. Biomechanical models and simulations can illuminate how extinct animals may have moved, but require anatomical data as inputs. With a phylogenetic context, osteological evidence, and neontological data on anatomy, it is possible to infer the musculature of extinct taxa. Here, we reconstructed the hindlimb musculature of Piatnitzkysauridae ( Condorraptor , Marshosaurus , and Piatnitzkysaurus ). We chose this clade for future usage in biomechanics, for comparisons with myological reconstructions of other theropods, and for the resulting evolutionary implications of our reconstructions; differential preservation affects these inferences, so we discuss these issues as well. We considered 32 muscles in total: for Piatnitzkysaurus , the attachments of 29 muscles could be inferred based on the osteological correlates; meanwhile, in Condorraptor and Marshosaurus , we respectively inferred 21 and 12 muscles. We found great anatomical similarity within Piatnitzkysauridae, but differences such as the origin of M . ambiens and size of M . caudofemoralis brevis are ev. Similarities were evident with Aves, such as the division of the M . iliofemoralis externus and M . iliotrochantericus caudalis and a broad depression for the M . gastrocnemius pars medialis origin on the cnemial crest. Nevertheless, we infer plesiomorphic features such as the origins of M . puboischiofemoralis internus 1 around the “cuppedicus” fossa and M . ischiotrochantericus medially on the ischium. As the first attempt to reconstruct muscles in early tetanurans, our study allows a more complete understanding of myological evolution in theropod pelvic appendages.
We report theropod caudal vertebrae found at Phu Wiang Mountain, Thailand. They resemble the Portuguese Baryonyx and pertain to the Spinosauridae based on the presence of striations on the surface of the transverse process of the caudals, well developed double keels and a deep ventral groove on the centra, two laminae, delimiting three fossae below the transverse process, and posterior caudals having curved, rod-like neural spines with small process at the base. This supports the presence of spinosaurids in the Sao Khua Formation of Thailand. Furthermore, the putative basal ceratosaur Camarillasaurus Sánchez- Hernández and Benton, 2014 from Spain is found here to be a spinosaurid based on the resemblance of the type materials to the Thai spinosaurid described in the present work, as well as the phylogenetic analysis and the similarity to other spinosaurids. In Europe, spinosaurids have been reported from England, Portugal, and Spain. The reassessment of Camarillasaurus adds to the number of this group in this region. The presence of more than one spinosaurid taxon in the same region is common and can be found in the Kem Kem Beds of Morocco, the Araripe Basin of Brazil, the Iberian Peninsula, and the Khorat Plateau of Southeast Asia.
Abelisaurid theropods dominated the predator role across Gondwana during the Late Cretaceous. They are characterized by highly reduced forelimbs and one of the most specialized cranial morphologies among carnivorous dinosaurs, exemplified by a broad skull, short rostrum, high occipital region, and highly kinetic intramandibular joint, suggestive of a specialized feeding strategy. Late Cretaceous abelisaurids are known from some remarkably complete taxa with well-preserved skulls. However, little is known about the pattern of character transformation that led to their highly modified condition because there are no well-preserved abelisaurids before the Late Cretaceous. Here we report a basal abelisaurid from the Early Cretaceous of Brazil that preserves a complete skull and reveals an early stage in the cranial evolution of the group. It lacks the specialized temporal and mandibular features characteristic of derived abelisaurids, including the kinetic intramandibular joint and knob-like dorsal projection of the parietals.
El presente trabajo analiza las variaciones morfológicas en huellas atribuidas a Bressaichnus patagonichus preservadas en la Formación Candeleros (Cañadón de las Campanas y Balneario el Chocón, Neuquén). El material comprende numerosas huellas de dinosaurios terópodos no-avianos incluidas en varias rastrilladas de las cuales se consideró el largo (FL) y ancho (FW) de la huella, el ángulo de divergencia total, y los ángulos entre los dígitos II-III y III-IV. Hay variación en aspectos formacionales y post-formacionales, producto de viento y agua. Los aspectos formacionales son postura (digitígrada y semidigitígrada) y sustrato (firmes y blandos). La evaluación de la variación se realizó a través de un análisis de componentes principales (PCA). En este, la primer componente (PC1) se ve mayormente influenciada por los ángulos interdigitales, y la segunda componente (PC2) por las variables dimensionales de largo y ancho de las huellas. En cuanto a la postura se observa una separación de digitígrados y semidigitígrados en PC1. Si bien, la definición morfológica aumenta con la pérdida de humedad, la preservación de huellas en sustratos blandos o firmes no influencia sustancialmente a la morfología de las huellas. El desgaste (ponderado de 0-3) tampoco influencia la morfología de las huellas; solo aquellas huellas que han sufrido el mayor desgaste (3), son diferenciables por su gran tamaño debido al oleaje y mareas. Así, se puede decir que la variabilidad observada se debe efectivamente al comportamiento del productor, y no así a otras variables tafonómicas o preservacionales.
Noasaurines form an enigmatic group of small-bodied predatory theropod dinosaurs known from the Late Cretaceous of Gondwana. They are relatively rare, with notable records in Argentina and Madagascar, and possible remains reported for Brazil, India, and continental Africa. In south-central Brazil, the deposits of the Bauru Basin have yielded a rich tetrapod fauna, which is concentrated in the Bauru Group. The mainly aeolian deposits of the Caiuá Group, on the contrary, bear a scarce fossil record composed only of lizards, turtles, and pterosaurs. Here, we describe the first dinosaur of the Caiuá Group, which also represents the best-preserved theropod of the entire Bauru Basin known to date. The recovered skeletal parts (vertebrae, girdles, limbs, and scarce cranial elements) show that the new taxon was just over 1 m long, with a unique anatomy among theropods. The shafts of its metatarsals II and IV are very lateromedially compressed, as are the blade-like ungual phalanges of the respective digits. This implies that the new taxon could have been functionally monodactyl, with a main central weight-bearing digit, flanked by neighbouring elements positioned very close to digit III or even held free of the ground. Such anatomical adaptation is formerly unrecorded among archosaurs, but has been previously inferred from footprints of the same stratigraphic unit that yielded the new dinosaur. A phylogenetic analysis nests the new taxon within the Noasaurinae clade, which is unresolved because of the multiple alternative positions that Noasaurus leali can acquire in the optimal trees. The exclusion of the latter form results in positioning the new dinosaur as the sister-taxon of the Argentinean Velocisaurus unicus.
The homology of the tridactyl hand of birds is a still debated subject, with both paleontological and developmental evidence used in support of alternative identity patterns in the avian fingers. With its simplified phalangeal morphology, the Late Jurassic ceratosaurian Limusaurus has been argued to support a II-III-IV digital identity in birds and a complex pattern of homeotic transformations in three-fingered (tetanuran) theropods. We report a new large-bodied theropod, Saltriovenator zanellai gen. et sp. nov., based on a partial skeleton from the marine Saltrio Formation (Sinemurian, lowermost Jurassic) of Lombardy (Northern Italy). Taphonomical analyses show bone bioerosion by marine invertebrates (first record for dinosaurian remains) and suggest a complex history for the carcass before being deposited on a well-oxygenated and well-illuminated sea bottom. Saltriovenator shows a mosaic of features seen in four-fingered theropods and in basal tetanurans. Phylogenetic analysis supports sister taxon relationships between the new Italian theropod and the younger Early Jurassic Berberosaurus from Morocco, in a lineage which is the basalmost of Ceratosauria. Compared to the atrophied hand of later members of Ceratosauria, Saltriovenator demonstrates that a fully functional hand, well-adapted for struggling and grasping, was primitively present in ceratosaurians. Ancestral state reconstruction along the avian stem supports 2-3-4-1-X and 2-3-4-0-X as the manual phalangeal formulae at the roots of Ceratosauria and Tetanurae, confirming the I-II-III pattern in the homology of the avian fingers. Accordingly, the peculiar hand of Limusaurus represents a derived condition restricted to late-diverging ceratosaurians and cannot help in elucidating the origin of the three-fingered condition of tetanurans. The evolution of the tridactyl hand of birds is explained by step-wise lateral simplification among non-tetanuran theropod dinosaurs, followed by a single primary axis shift from digit position 4 to 3 at the root of Tetanurae once the fourth finger was completely lost, which allowed independent losses of the vestigial fourth metacarpal among allosaurians, tyrannosauroids, and maniraptoromorphs. With an estimated body length of 7.5 m, Saltriovenator is the largest and most robust theropod from the Early Jurassic, pre-dating the occurrence in theropods of a body mass approaching 1,000 Kg by over 25 My. The radiation of larger and relatively stockier averostran theropods earlier than previously known may represent one of the factors that ignited the trend toward gigantism in Early Jurassic sauropods.
Ceratosaur theropods ruled the Southern Hemisphere until the end of the Late Cretaceous. However, their origin was earlier, during the Early Jurassic, a fact which allowed the group to reach great morphological diversity. The body plans of the two main branches (Noasauridae and new name Etrigansauria: Ceratosauridae + Abelisauridae) are quite different; nevertheless, they are sister taxa. Abelisaurids have lost the ability to grasp in the most derived taxa, but the reduced forelimb might have had some display function. The ontogenetic changes are well known in Limusaurus which lost all their teeth and probably changed the dietary preference at maturity. The results presented here suggest that abelisaurids had different soft tissues on the skull. These tissues might have been associated with evolution of a strong cervicocephalic complex and should have allowed derived taxa (e.g.
Majungasaurus and Carnotaurus) to have low-displacement headbutting matches. The ability to live in different semi-arid environment plus high morphological disparity allowed the ceratosaurs to become an evolutionary success.
Dinosauromorphs evolved a wide diversity of hind limb skeletal morphologies, suggesting highly divergent articular soft tissue anatomies. However, poor preservation of articular soft tissues in fossils has hampered any follow-on functional inferences. We reconstruct the hip joint soft tissue anatomy of non-dinosaurian dinosauromorphs and early dinosaurs using osteological correlates derived from extant sauropsids and infer trends in character transitions along the theropod and sauropodomorph lineagues. Femora and pelves of 107 dinosauromorphs and outgroup taxa were digitized using 3D imaging techniques. Key transitions were estimated using maximum likelihood ancestral state reconstruction. The hips of dinosauromorphs possessed wide a disparity of soft tissue morphologies beyond the types and combinations exhibited by extant archosaurs. Early evolution of the dinosauriform hip joint was characterized by the retention of a prominent femoral hyaline cartilage cone in post-neonatal individuals, with the cartilage cone independently reduced within theropods and sauropodomorphs. The femur of Dinosauriformes possessed a fibrocartilage sleeve on the metaphysis, which surrounded a hyaline core. The acetabulum of Dinosauriformes possessed distinct labrum and antitrochanter structures. In sauropodomorphs, hip congruence was maintained by thick hyaline cartilage on the femoral head, whereas theropods relied on acetabular tissues such as ligaments and articular pads. In particular, the craniolaterally ossified hip capsule of non-Avetheropoda neotheropods permitted mostly parasagittal femoral movements. These data indicate that the dinosauromorph hip underwent mosaic evolution within the saurischian lineage and that sauropodomorphs and theropods underwent both convergence and divergence in articular soft tissues, correlated with transitions in body size, locomotor posture, and joint loading.
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Citation for this article: Tsai, H. P., K. M. Middleton, J. R. Hutchinson, and C. M. Holliday. 2018. Hip joint articular soft tissues of non-dinosaurian Dinosauromorpha and early Dinosauria: evolutionary and biomechanical implications for Saurischia. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1427593.
During the latest Cretaceous, distinct dinosaur faunas were found in Laurasia and Gondwana. Tyrannosaurids, hadrosaurids, and ceratopsians dominated in North America and Asia, while abelisaurids and titanosaurids dominated in South America, India, and Madagascar. Little is known about dinosaur faunas from the latest Cretaceous of Africa, however. Here, a new abelisaurid theropod, Chenanisaurus barbaricus, is described from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa on the basis of a partial dentary and isolated teeth. Chenanisaurus is both one of the largest abelisaurids, and one of the youngest known African dinosaurs. Along with previously reported titanosaurid remains, Chenanisaurus documents the persistence of a classic Gondwanan abelisaurid-titanosaurid fauna in mainland Africa until just prior to the end-Cretaceous mass extinction. The animal is unusual both in terms of its large size and the unusually short and robust jaw. Although it resembles South American carnotaurines in having a deep, bowed mandible, phylogenetic analysis suggests that Chenanisaurus may represent a lineage of abelisaurids that is distinct from those previously described from the latest Cretaceous of South America, Indo-Madagascar, and Europe, consistent with the hypothesis that the fragmentation of Gondwana led to the evolution of endemic dinosaur faunas during the Late Cretaceous.
Abelisauroids are the most abundant theropods in the Cretaceous beds of Patagonia. They are traditionally subdivided into large-sized Abelisauridae and smaller Noasauridae. Here, we describe a new specimen of the small enigmatic abelisauroid Velocisaurus unicus Bonaparte, 1991, which was previously known from a single incomplete specimen from Neuquén City, Neuquén Province, Patagonia. The new material comes from the Santonian Bajo de la Carpa Formation (Late Cretaceous) at the Paso Córdova locality, Río Negro Province. It comprises an almost complete left hind limb and offers novel information about the anatomy of this poorly known abelisauroid. The new material shows that Velocisaurus is remarkable in having a very short, stout, and anteriorly bowed femur, which has a notably subtriangular cross-section at its proximal end. The tibia is long and slender, and the anterior surface of the distal end is anteroposteriorly flat and transversely expanded, with an enlarged surface for the ascending process of the astragalus. The pes has a stout third metatarsal, rod-like metatarsals II and IV, and highly modified phalanges of digit IV. The unique combination of characters of Velocisaurus indicates that this taxon belongs to a still poorly understood radiation of gracile-limbed abelisauroids. The inclusion of Velocisaurus in a comprehensive phylogenetic analysis recovers a monophyletic Noasauridae, but with only very weak support. Detailed analysis of features supporting the inclusion of Velocisaurus within Noasauridae is discussed, and their implications for abelisauroid phylogeny are revisited. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Brissón Egli, F., F. L. Agnolín, and Fernando Novas. 2016. A new specimen of Velocisaurus unicus (Theropoda, Abelisauroidea) from the Paso Córdoba locality (Santonian), Río Negro, Argentina. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1119156.
Many recent studies of theropod relationships have been focused on the phylogeny of coelurosaurs and the question of the origin of birds, but the interrelationships and evolution of basal theropods are still poorly understood. Thus, this paper presents a phylogenetic analysis of all theropods, but focuses on the basal members of this clade. The result supports the inclusion of Eoraptor and herrerasaurids in the Theropoda, but differs from other recent studies in two main aspects: (1) The taxa usually grouped as ceratosaurs form two monophyletic clades that represent successively closer outgroups to tetanurans. The more basal of these clades, the Coelophysoidea, comprise the majority of Late Triassic and Early Jurassic theropods. The other clade of basal theropods that are usually included in the Ceratosauria comprises Ceratosaurus, Elaphrosaurus, and abelisaurids. (2) Two monophyletic groups of basal tetanurans are recognized: the Spinosauroidea and the Allosauroidea. In contrast to other recent phylogenetic hypotheses, both clades are united in a monophyletic Carnosauria. The branching pattern of the present cladogram is in general accordance with the stratigraphic occurrence of theropod taxa. Despite the differences in recent analyses, there is a significant level of consensus in theropod phylogeny. At least four different radiations of non-avian theropods can be recognized. These radiations show different patterns in Laurasia and Gondwana, and there are increasing differences between the theropod faunas of the two hemispheres from the Triassic to the Cretaceous.
Resumen
The Late Cretaceous (Maastrichtian) Lameta Formation of central India has yielded dissociated elements of a variety of predatory dinosaurs, most of them coming from a quarry named the "Carnosaur bed." The materials were described by Huene and Matley nearly 70 years ago. They recognized nine theropod species, which they sorted out into the theropod subgroups "Carnosauria" and "Coelurosauria". Huene and Matley also described a considerable amount of theropod hindlimb bones (e.g., femora, tibiae, metatarsals, and pedal phalanges) that they could not refer to any of these species, but vaguely interpreted as corresponding to "allosaurid" or "coelurosaurid" theropods. We reviewed the available collection of Cretaceous theropods from Bara Simla housed at the Geological Survey of India, Calcutta, arriving to the following conclusions: 1) Indosuchus and Indosaurus are abelisaurids, as recognized by previous authors, but available information is not enough to judge whether they are synonyms; 2) Laevisuchus indicus is a small abelisauroid, related to Noasaurus and Masiakasaurus on the basis of their peculiar cervical vertebrae; 3) the controversial taxa " Compsosuchus", " Dryptosauroides", " Ornithomimoides", and " Jubbulpuria" are represented by isolated vertebrae corresponding to different portions of the neck and tail, and also exhibit abelisauroid features; 4) hindlimb bones originally referred to as "allosaurid" and "coelurosaurian" also exhibit abelisauroid characters, and bones of large size are tentatively referred to as corresponding to Indosuchus or Indosaurus, whereas some pedal bones of smaller size may belong to Laevisuchus; 5) two kinds of abelisaurid feet are apparent: one in which the phalanges of digit III and IV are robust, and another type in which the phalanges of digit IV are transversely narrow and dorsoventrally deep. This review demonstrates that all of the theropod elements discovered at the "Carnosaur bed" belong to a single theropod clade, the Abelisauroidea
Living crocodilians and limbed lepidosaurs have a large caudofemoralis longus muscle passing from tail to femur. Anatomical and electromyographic data support the conclusion that the caudofemoralis is the principal femoral retractor and thus serves as the primary propulsive muscle of the hind limb. Osteological evidence of both origin and insertion indicates that a substantial caudofemoralis longus was present in archosaurs primitively and was retained in the clades Dinosauria and Theropoda. Derived theropods (e.g., ornithomimids, deinonychosaurs, Archaeopteryx and birds) exhibit features that indicate a reduction in caudofemoral musculature, including fewer caudal vertebrae, diminished caudal transverse processes, distal specialization of the tail, and loss of the fourth trochanter. This trend culminates in ornithurine birds, which have greatly reduced tails and either have a minute caudofemoralis longus or lack the muscle entirely.
As derived theropod dinosaurs, birds represent the best living model for reconstructing extinct nonavian theropods. Bipedal, digitigrade locomotion on fully erect limbs is an avian feature inherited from theropod ancestors. However, the primitive saurian mechanisms of balancing the body (with a large tail) and retracting the limb (with the caudofemoralis longus) were abandoned in the course of avian evolution. This strongly suggests that details of the orientation (subhorizontal femur) and movement (primarily knee flexion) of the hind limb in extant birds are more properly viewed as derived, uniquely avian conditions, rather than as retentions of an ancestral dinosaurian pattern. Although many characters often associated with extant birds appeared much earlier in theropod evolution, reconstructing the locomotion of all theropods as completely birdlike ignores a wealth of differences that characterize birds.
Ceratosauria represents the first widespread and diverse radiation of theropod dinosaurs comprising two main sister clades, Neoceratosauria and Coelophysoidea. This chapter discusses the diagnostic features, phylogenetic placement, and paleobiology of ceratosaurians. The fossil record for Ceratosauria spans a minimum of 155 million years, from the late Carnian of the Late Triassic to the end of the Cretaceous. Ceratosaurs had an essentially global distribution, their remains being found on every continent except Australia and Antarctica. Ceratosaurs evolved into a broad range of sizes and body forms, from lightly built, diminutive taxa such as Segisaurus (1 to 1.5 m in length) to the large abelisaurids, such as Carnotaurus (10 to 11 m). Several coelophysoid taxa were collected from mass burials, where multiple individuals were preserved together. In particular, Syntarsus rhodesiensis is known from at least thirty individuals found at localities in Zimbabwe and South Africa.
In this article, we develop a new reconstruction of the pelvic and hindlimb muscles of the large theropod dinosaur Tyrannosaurus rex. Our new reconstruction relies primarily on direct examination of both extant and fossil turtles, lepidosaurs, and archosaurs. These observations are placed into a phylogenetic context and data from extant taxa are used to constrain inferences concerning the soft-tissue structures in T. rex. Using this extant phylogenetic bracket, we are able to offer well-supported inferences concerning most of the hindlimb musculature in this taxon. We also refrain from making any inferences for certain muscles where the resulting optimizations are ambiguous. This reconstruction differs from several previous attempts and we evaluate these discrepancies. In addition to providing a new and more detailed understanding of the hindlimb morphology of T. rex--the largest known terrestrial biped--this reconstruction also helps to clarify the sequence of character-state change along the line to extant birds.
Abelisaurids are a clade of large, bizarre predatory dinosaurs, most notable for their high, short skulls and extremely reduced forelimbs. They were common in Gondwana during the Cretaceous, but exceedingly rare in the Northern Hemisphere. The oldest definitive abelisaurids so far come from the late Early Cretaceous of South America and Africa, and the early evolutionary history of the clade is still poorly known. Here, we report a new abelisaurid from the Middle Jurassic of Patagonia, Eoabelisaurus mefi gen. et sp. nov., which predates the so far oldest known secure member of this lineage by more than 40 Myr. The almost complete skeleton reveals the earliest evolutionary stages of the distinctive features of abelisaurids, such as the modification of the forelimb, which started with a reduction of the distal elements. The find underlines the explosive radiation of theropod dinosaurs in the Middle Jurassic and indicates an unexpected diversity of ceratosaurs at that time. The apparent endemism of abelisauroids to southern Gondwana during Pangean times might be due to the presence of a large, central Gondwanan desert. This indicates that, apart from continent-scale geography, aspects such as regional geography and climate are important to reconstruct the biogeographical history of Mesozoic vertebrates.
The Late Cretaceous Marília Formation (Bauru Group, Bauru Basin) is a geological unit that occurs on São Paulo, Minas Gerais, Goiás, and Mato Grosso do Sul states, Brazil. This formation consists predominantly of paleosols developed in a semiarid/arid environment and recent reappraisal of its formerly known members reduced its lithological composition and geographical distribution. Hence, the Marília Formation has a very sparse vertebrate fossil record without named species so far. In this contribution we describe a new abelisaurid theropod (Dinosauria) from this unit, namely Kurupi itaata gen. et sp. nov., discovery in the Municipality of Monte Alto, western São Paulo State. The holotype MPMA 27-0001/02 consists of three caudal vertebrae and the partial pelvic girdle. Kurupi itaata gen. et sp. nov. shares with other South American abelisaurids fused ischia and caudal vertebrae with long and laterodorsally oriented transverse processes, with fan-shaped distal ends. Autapomorphies of the new theropod include the variation of ∼15º of the inclination of the transverse process between the first (C1) and seventh (C7) caudal vertebra; C1 with anterolaterally projected triangular process placed below the proximodistal mid-portion of the transverse process, and a notch at the anterodistal portion of the transverse process, between a shelf-like process and the anterodistal corner of the transverse process; and anterior caudal vertebrae (present in C1 and C7, inferred in the others) with a cuneiform process, anterodorsally projected, located on the dorsal surface of the transverse process. Phylogenetic analysis recovered Kurupi itaata gen. et sp. nov. among abelisaurid theropods, but was nested in an unresolved massive polytomy of the entire clade. Taphonomic traits on the studied specimens corroborate previous proposals for the paleoenvironmental context of the Marília Formation. Kurupi itaata gen. et sp. nov. was about 5 meters long, with a rigid tail, and cursorial locomotion as indicated by its a muscles attachment and bones anatomy. This new taxon contributes to the knowledge of the Maastrichtian continental fauna of Brazil and increases the diversity of medium-sized abelisaurids in western Gondwana.
Abelisauridae is a theropod clade with a wide distribution in the Late Cretaceous of Gondwana. Some of the best preserved abelisaurid specimens were recovered from Patagonia (Argentina) such as Skorpiovenator, Ilokelesia, Carnotaurus and Aucasaurus. Here we describe a dorsal part of a neural spine; a middle caudal vertebra; a distal part of a left metatarsal IV; a complete right phalanx IV-1; left phalanges IV-1, 2 and 3; and a pedal ungual phalanx. These materials were recovered from the same quarry of a recently published indeterminate abelisaurid specimen (MPCN-PV-69). The most distinctive characters are a triangular shape of the distal end of metatarsal IV (present in other abelisauroids); phalanx IV-1 with the proximal surface dorsoventrally tall and the ventral surface wider than the dorsal one causing a medial tilting of bone (set of features considered autapomorphic in Velocisaurus, although is also present in different abelisaurids); phalanx IV-1 and 2 with a ridge which spans from the proximodorsal projection, splits in two branch and surrounds a laterally displaced and obliquely oriented oval hyperextensor pit; a pedal ungual phalanx with two medial and lateral vascular grooves, and lacking a flexor tubercle (abelisauroids synapomorphies). The mentioned feature of phalanges IV-1 and 2 is only found among abelisaurids and is here considered as a possible new synapomorphy of Abelisauridae. Thus, besides they constitute new abelisaurid remains from Patagonia, the new materials provide valuable morphological data that could expand the diagnosis of Abelisauridae.
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.
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.
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.
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.
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.
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).
A new genus and species of ornithomimosaur, Afromimus tenerensis, is described based on a fragmentary skeleton from the Lower Cretaceous (Aptian—Albian) El Rhaz Formation of Niger. The holotype and only known individual preserves caudal vertebrae, chevrons and portions of the right hind limb. Derived ornithomimosaurian features include the broad, peanut-shaped articular surfaces of mid caudal centra, parasagittal fossae on mid caudal centra for reception of the postzygapophyses of the preceding vertebra, and a raised, subtriangular platform on the ventral aspect of the pedal phalanges. New information is given for, and comparisons made to, Nqwebasaurus thwazi from southern Africa, the oldest and most basal ornithomimosaur. Unlike other coelurosaurian clades that have expansive radiations on northern landmasses, the oldest ornithomimosaur and now another basal form are known from a southern landmass, Africa.
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.
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).
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.
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.
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.
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.
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.
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.
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.
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.
