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Osteology of Aerosteon riocoloradensis (Sereno et al. 2008) a large megaraptoran (Dinosauria: Theropoda) from the Upper Cretaceous of Argentina

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Abstract

Aerosteon riocoloradensis represents one of the most complete megaraptorans yet discovered. This theropod comes from Anacleto Formation (Campanian) of Mendoza Province, Argentina. The aims of this contribution are: to present a detailed, bone by bone description of this specimen with figures of each bone; provide comparisons to other closely related theropods; revise the original assignation and diagnosis of such taxa. Three bones were re-assigned and almost all the autapomorphies of Aerosteon were modified. Features in the vertebral columns, which are shared with other megaraptorans, show that these theropods shared features with basal coelurosaurs. Anatomical Abbreviations ACDL: Anterior centrodiapophyseal lamina; CDF: Centrodiapophyseal fossa; CPAL: Centroparapophyseal lamina; CPRL: Centroprezygapophyseal lamina; CPRF: Centroprezygapophyseal fossa; CPR-CDF: Centroprezygapophyseal-centrodiapophyseal fossa; Hye: Hyposphene; Hym: Hypantrum; ILT: Intervertebral ligament tuberosity; IPOL: Infrapostzygapophysela lamina; IZL: Intrazygapophyseal lamina; PADL: Paradiapophyseal lamina; PAD-CDF: Paradiapophyseal-centrodiapophsyeal fossa; PCDL: Posterior centrodiapophyseal lamina; POEL: Postzygaepipophysela lamina; PODL: Postzygadiapophyseal lamina; POSF: Postspinal fossa; POCDF: Postzygapophsyeal-centrodiapophyseal fossa; Poz: Postzygapophysis; PRDL: Prezygadiapophyseal lamina; PRPAF: Prezygaparapophyseal fossa; PRPAL: Prezygaparapophyseal lamina; PRSF: Prespinal fossa; PRSL: Prespinal lamina; PRD-CDF: Prezygadiapophyseal-centrodiapophyseal fossa; PRD-PADF: Prezygadiapophyseal-paradiapophyseal fossa; PRD-PODF: Prezygadiapophyseal-postzygadiapophyseal fossa; PRCDF: Prezygapophyseal-centrodiapophyseal fossa; Prz: Prezygapophyses; SDF: Supradiapophsyeal fossa; SDL: Supradiapophyseal lamina; SPOF: Spinopostzygapophyseal fossa; SPOL: Spinopostzygapophyseal lamina; SPRF: Spinoprezygapophyseal fossa; SPRL: Spinoprezygapophyseal lamina; SR(number): Sacral rib; STP(number): Sacral transverse process

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... Only three caudal vertebrae are currently known for Maip macrothorax, and all are from the proximal third of the tail, thereby enabling comparison with the proximal caudals of Megaraptor namunhuaiquii 10 , Orkoraptor burkei 35 , and Aerosteon riocoloradensis 14,34 . The holotype of Maip macrothorax also preserves several ribs tentatively referred as cervical ribs 5, 7 and 8 (Fig. 10), and dorsal ribs 1, 2, and 6, as well as many other indeterminate rib fragments (Fig. 11). ...
... Dorsal 4. It is represented by an isolated centrum ( Fig. 4F-I). This element is identified as the fourth because part of the parapophysis is placed in the neural arch, a single squared or round pneumatopore, as well as the absence of a ventral keel, a condition retained in the first dorsal elements of theropods (such as D1 of Aerosteon; 14,34 ). Furthermore, this interpretation is tentative giving the lack of articulated dorsal columns in non-juvenile megaraptorid specimens. ...
... Interestingly, the prezygapophyses of Maip are placed below in the neural arch and do not reach the base of transverse process and the base of the neural spine. This condition is shared with the D4 of Aerosteon and the first dorsal vertebrae of derived tyrannosaurids 14,44 . In contrasts, the prezygapophyses surpass the dorsal margin of the base of the transverse process in D3 of the mid-sized megaraptorid Murusraptor, and the D2 of small-sized tyrannosauroids (such as Guanlong 58 ), as well as most tetanurans such as Allosaurus, Sinraptor, Lajasvenator and Neovenator 43,45,59,60 . ...
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The quadrate of reptiles and most other tetrapods plays an important morphofunctional role by allowing the articulation of the mandible with the cranium. In Theropoda, the morphology of the quadrate is particularly complex and varies importantly among different clades of non-avian theropods, therefore conferring a strong taxonomic potential. Inconsistencies in the notation and terminology used in discussions of the theropod quadrate anatomy have been noticed, including at least one instance when no less than eight different terms were given to the same structure. A standardized list of terms and notations for each quadrate anatomical entity is proposed here, with the goal of facilitating future descriptions of this important cranial bone. In addition, an overview of the literature on quadrate function and pneumaticity in non-avian theropods is presented, along with a discussion of the inferences that could be made from this research. Specifically, the quadrate of the large majority of non-avian theropods is akinetic but the diagonally oriented intercondylar sulcus of the mandibular articulation allowed both rami of the mandible to move laterally when opening the mouth in many of theropods. Pneumaticity of the quadrate is also present in most averostran clades and the pneumatic chamber—invaded by the quadrate diverticulum of the mandibular arch pneumatic system—was connected to one or several pneumatic foramina on the medial, lateral, posterior, anterior or ventral sides of the quadrate.
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Megaraptoridae comprises a clade of enigmatic Gondwanan theropods with characteristic hypertrophied claws on the first and second manual digits. The majority of megaraptorids are known from South America, although a single genus (Australovenator) plus additional indeterminate material is also known from Australia. This clade has a controversial placement among theropods, and recently has been interpreted alternatively as a carcharodontosaurian or a tyrannosauroid lineage. We describe new fragmentary but associated postcranial remains from the opal fields of Lightning Ridge (middle-Albian, Griman Creek Formation) in north-central New South Wales. The new unnamed taxon exhibits a number of unusual features that suggest the presence of a hitherto unrecognized Australian megaraptorid. From an Australian perspective, the Lightning Ridge taxon predates Australovenator by c.10Ma and is minimally coeval with megaraptoran material reported from the Eumeralla Formation of Victoria (but potentially 6.1–9.5 Ma younger). It is also notable as the largest predatory dinosaur yet identified from Australia and is only the second theropod known from more than a single element. A Bayesian phylogenetic approach integrating morphological, stratigraphic and palaeogeographic information tested both the carcharodontosaurian and tyrannosauroid placements for Megaraptora. Regardless of the preferred placement among Tetanurae, rigorous palaeobiogeographic analyses support an Asian origin of Megaraptora in the latest Jurassic (about 150-135 Ma), an Early Cretaceous (about 130-121 Ma) divergence of the Gondwanan lineage leading to Megaraptoridae, and an Australian root for megaraptorid radiation. These results indicate that Australia’s Cretaceous dinosaur fauna did not comprise simply of immigrant taxa but was a source for complex two-way interchange between Australia-Antarctica-South America leading to the evolution of at least one group of apex predatory dinosaurs in Gondwana.
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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.
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Allosauroid theropods were a diverse and widespread radiation of Jurassic-Cretaceous megapredators. Achieving some of the largest body sizes among theropod dinosaurs, these colossal hunters dominated terrestrial ecosystems until a faunal turnover redefined apex predator guild occupancy during the final 20 million years of the Cretaceous. Here we describe a giant new species of allosauroid - Siats meekerorum gen. et sp. nov. - providing the first evidence for the cosmopolitan clade Neovenatoridae in North America. Siats is the youngest allosauroid yet discovered from the continent and demonstrates that the clade endured there into the Late Cretaceous. The discovery provides new evidence for ecologic sympatry of large allosauroids and small-bodied tyrannosauroids. These data support the hypothesis that extinction of Allosauroidea in terrestrial ecosystems of North America permitted ecological release of tyrannosauroids, which went on to dominate end-Cretaceous food webs.
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We performed additional preparation on the holotype skeleton of Nqwebasaurus thwazi and discovered new skeletal material. We describe this material, which includes a maxilla with small, conical, unserrated teeth and bones of the braincase, as well as parts of the holotype postcranial anatomy that were previously poorly documented. We incorporate this new anatomical information into a broadly sampled matrix designed to test theropod relationships. Our phylogenetic results hypothesize that Nqwebasaurus is the basalmost ornithomimosaur, and recover numerous characters supporting this relationship, including features of the maxilla, frontal, dentition, axial skeleton, forelimb and hindlimb. Nqwebasaurus is the first African ornithomimosaur and the first Gondwanan member of this group known from articulated skeletal material, supporting the hypothesis that coelurosaurian groups were cosmopolitan during their early evolutionary history. The presence of reduced dentition and a gastric mill in Nqwebasaurus strongly suggest that this taxon was herbivorous.
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We report new skeletal elements pertaining to the same individual which represents the holotype of Australovenator wintonensis, from the 'Matilda Site' in the Winton Formation (Upper Cretaceous) of western Queensland. The discovery of these new elements means that the hind limb of Australovenator is now the most completely understood hind limb among Neovenatoridae. The new hind limb elements include: the left fibula; left metatarsal IV; left pedal phalanges I-2, II-1, III-4, IV-2, IV-3; and right pedal phalanges, II-2 and III-1. The detailed descriptions are supported with three dimensional figures. These coupled with the completeness of the hind limb will increase the utility of Australovenator in comparisons with less complete neovenatorid genera. These specimens and the previously described hind limb elements of Australovenator are compared with other theropods classified as neovenatorids (including Neovenator, Chilantaisaurus, Fukuiraptor, Orkoraptor and Megaraptor). Hind limb length proportion comparisons indicate that the smaller neovenatorids Australovenator and Fukuiraptor possess more elongate and gracile hind limb elements than the larger Neovenator and Chilantaisaurus. Greater stride lengths to body size exist in both Fukuiraptor and Australovenator with the femur discovered to be proportionally shorter the rest of the hind limb length. Additionally Australovenator is identified as possessing the most elongate metatarsus. The metatarsus morphology varies with body size. The larger neoventorids possess a metatarsus with greater width but shorter length compared to smaller forms.
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We describe a new coelurosaurian theropod, Zuolong salleei, gen. et sp. nov., from exposures of the upper part of the Shishugou Formation at the Wucaiwan locality, Xinjiang Autonomous Region, People's Republic of China. Zuolong has a large, inclined quadrate foramen that extends onto the medial surface of the quadratojugal, an unusually large fovea capitis on the femoral head, and an apomorphically large distal condyle of metatarsal III with a medially projecting flange on the extensor surface. Radiometric dating of the Shishugou Formation constrains the age of the specimen to the beginning of the Late Jurassic (Oxfordian). A cladistic analysis of Zuolong salleei in a broadly sampled theropod data matrix recovers it as a basal coelurosaur. These data make Zuolong one of the oldest coelurosaur fossils yet known that preserves both cranial and postcranial bones.
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Therizinosauria are an unusual group of theropod dinosaurs, found mostly in the Cretaceous deposits in Mongolia, China and western USA. The basal forms of this group are represented by incomplete or disarticulated material. Here, we report a nearly complete, articulated skeleton of a new basal therizinosaur from the Early Cretaceous Yixian Formation of Jianchang County, western part of Liaoning Province, which sheds light on our understanding of anatomy of basal therizinosaurs. This new dinosaur shows some typical therizinosaur features, such as neural spines of the anterior caudal vertebrae that possess anterior and posterior alae, a rectangular buttress on the ventrolateral side of the proximal end of metacarpal I, and appressed metatarsal shafts. Our phylogenetic analysis suggests that it is a basal therizinosaur (sister taxon to Therizinosauroidea) because it bears many basal therizinosaur characters in the dentition, pelvis and hind limbs. The new therizinosaur described here has unique tooth and jaw characters such as the offsetting of the tooth row by a shelf and dentary teeth with labially concave and lingually convex dentary teeth, similar to ornithopods and ceratopsians.
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The theropod dinosaur Aniksosaurus darwini gen. et sp. nov. has been recovered from the Upper Cretaceous, Bajo Barreal Formation, of Central Patagonia. Aniksosaurus darwini gen. et sp. nov. was a small tetanurine, approximately 2 meters long. Aniksosaurus exhibits several unique traits (e.g., cranial cervical ver- tebrae with dorsoventrally deep neural arches, provided with a pair of cavities at their cranial surfaces; neural canal wide; cranial caudals with ventral sagittal keel, and transverse processes triangular-shaped in dorsal view; manual ungual phalanges robust; ilium with extremely expanded brevis shelf; femur with deep notch for M. Iliotrochantericus; metatarsal and digit IV of pes transversely narrow). Available postcranial bones of Aniksosaurus exhibit derived features of Coelurosauria (e.g., ilium with well developed cuppedicus fossa; femur with anterior trochanter proximally projected, almost reaching the level of the articular head; greater tro- chanter craniocaudally expanded; femoral head rectangular-shaped in cranial aspect; and fibular shaft craniocaudally narrow), as well as characteristics suggesting that the new Patagonian taxon is more derived than some basal coelurosaurians such as compsognathids, Ornitholestes, and coelurids. Comparisons with maniraptoriforms (a clade including Ornithomimosauria, Tyrannosauridae, Oviraptorosauria, Alvarezsauridae and Paraves) support that Aniksosaurus is less derived than these theropods. In sum, Aniksosaurus is here considered as a Late Cretaceous survivor of a basal coelurosaurian radiation.
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Late Cretaceous (Cenomanian) fossils discovered in the Kem Kem region of Morocco include large predatory dinosaurs that inhabited Africa as it drifted into geographic isolation. One, represented by a skull approximately 1.6 meters in length, is an advanced allosauroid referable to the African genus Carcharodontosaurus. Another, represented by a partial skeleton with slender proportions, is a new basal coelurosaur closely resembling the Egyptian genus Bahariasaurus. Comparisons with Cretaceous theropods from other continents reveal a previously unrecognized global radiation of carcharodontosaurid predators. Substantial geographic differentiation of dinosaurian faunas in response to continental drift appears to have arisen abruptly at the beginning of the Late Cretaceous.
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A new ornithopod dinosaur, the medium-sized Sektensaurus sanjuanboscoi gen. et sp nov. from the Coniacian–Maastricthian strata of the Lago Colhué Huapi Formation, Golfo San Jorge Basin, Patagonia, is here described. The specimen consists of disarticulated postcranial bones belonging to a single individual. It is diagnosed by an unique association of five autapomorphies on the dorsal and sacral vertebrae and the humerus. A cladistics analysis indicates that Sektensaurus was an iguanodontian ornithopod which inhabited Patagonia during the Late Cretaceous. Likewise, Sektensaurus is the first non-hadrosaurid, probably elasmarian, ornithopod recorded from the uppermost Cretaceous of central Patagonia. In a broad context, this discovery increases the anatomical knowledge of ornithopods and adds new data on the composition of dinosaur faunas that lived in Patagonia close to the Antarctica, at the end of the Cretaceous.
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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).
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The mid-Cretaceous Griman Creek Formation (GCF), which crops out near the town of Lightning Ridge in the Surat Basin of north-central New South Wales, Australia, is noteworthy for its opalised vertebrate fauna. The fossil assemblage comprises remains of aspidorhynchid teleosts, lamniform chondrichthyans, dipnoans, chelid and possible meiolaniform turtles, leptocleidid-like and possible elasmosaurid plesiosaurians, anhanguerian pterosaurs, titanosauriform sauropods, megaraptoran theropods, ankylosaurians, several forms of non-iguanodontian and iguanodontian ornithopods, crocodylomorphs, enantiornithine birds, and stem and true monotremes, making it one of the most diverse mid-Cretaceous terrestrial vertebrate faunas in Australia. A detailed stratigraphic survey of twenty subterranean opal mines provides new information on the geology, age and palaeoenvironment of the main fossil-bearing beds. Vertebrate remains derive from the ‘Finch Clay facies’ laterally-extensive but discontinuous lenses of claystone that likely accumulated relatively rapidly in near-coastal but freshwater embayments (i.e. lagoonal conditions), and probably represent a single, roughly contemporaneous fauna. U-Pb age dating of detrital zircons extracted from a distinct layer of volcanogenic claystone immediately overlying one of the opalised fossil-bearing layers yields a maximum depositional age of 100.2–96.6 Ma. These new dates confirm an early to mid-Cenomanian age for the fauna, rather than Albian, as has been reported previously. The GCF at Lightning Ridge is therefore equivalent to the middle part of the Winton Formation (Queensland) and several million years older than the sauropod-dominated fauna at Winton.
Article
We describe Tratayenia rosalesi gen. et sp. nov., a new megaraptoran theropod dinosaur from the Upper Cretaceous of Patagonia, Argentina. The holotype consists of a well-preserved, mostly articulated series of dorsal and sacral vertebrae, two partial dorsal ribs, much of the right ilium, and pubis and ischium fragments. It was found in a horizon of the Upper Cretaceous (Santonian) Bajo de la Carpa Formation of the Neuquén Group in the Neuquén Basin exposed near the town of Añelo in Neuquén Province of northwestern Patagonia. Phylogenetic analysis recovers Tratayenia within the Gondwanan megaraptoran subclade Megaraptoridae. The new taxon exhibits similarities to other megaraptorids such as Aerosteon riocoloradensis, Megaraptor namunhuaiquii, and Murusraptor barrosaensis, but also presents differences in the architecture of the dorsal and sacral vertebrae and the morphology of the ilium. Tratayenia is the first megaraptoran that unequivocally preserves the complete sequence of sacral vertebrae, thereby increasing knowledge of the osteology of the clade. Moreover, depending on the chronostratigraphic ages of the stratigraphically controversial megaraptorids Aerosteon and Orkoraptor burkei, as well as the phylogenetic affinities of several fragmentary specimens, the new theropod may be the geologically youngest megaraptorid or megaraptoran yet discovered. Tratayenia is also the largest-bodied carnivorous tetrapod named from the Bajo de la Carpa Formation, reinforcing the hypothesis that megaraptorids were apex predators in southern South America from the Turonian through the Santonian or early Campanian, following the extinction of carcharodontosaurids.
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A specimen composed of a partial sacrum articulated to two anterior caudal vertebrae and an ilium (SMNS 58023) from the Lower Cretaceous (Albian) Santana Formation of Brazil was originally described as an oviraptorosaur, but it is here re-interpreted as the oldest megaraptoran of South America. The phylogenetic relationships of SMNS 58023 were tested quantitatively for the first time including it in the two most comprehensive phylogenetic data sets focused on non-maniraptoran theropods –including megaraptorans. The Brazilian specimen was consistently found as a megaraptoran in both analyses because of the presence of sacral centra longer than tall, absence of a median transverse constriction of sacral centra, and the morphology and position of sacral pleurocoels. SMNS 58023 sheds light on a region of the body that is poorly known in megaraptorans and pulls back the temporal range of the clade in South America. This re-interpretation reinforces the absence of oviraptorosaurs in Gondwana.
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A new compsognathid dinosaur, Sinocalliopteryx gigas gen. et sp. nov., is erected based on a complete skeleton from the Early Cretaceous Yixian Formation of western Liaoning, northeastern China. It shares the features with Huaxiagnathus orientalis in having a manus as long as the humerus plus radius, very large and subequally long manual claws I and II, and reduced olecranon process on the ulna. But it differs from Huaxiagnathus orientalis in having the much large size, a very long maxillary process of premaxilla not extending the vertical level of the maxillary antorbital fossa, and the proportionally longer ulna and so on. Sinocalliopteryx gigas gen. et sp. nov. represents the largest species among the known compsognathid dinosaurs, suggesting the tendency of the body enlargement in compsognathids to some extent. The long filamentous integuments are attached to the whole body of this compsognathid, confirming that such integuments evolved firstly in the basal coelurosaurs. This new giant compsognathid was a fierce carnivorous theropod, as shown further by an incomplete dromaeosaurid leg inside its abdominal cavity.
Chapter
Tetanurae comprised Carnosauria, a monophyletic group composed of Allosaurus, and Coelurosauria, containing modern birds and all taxa closer to modern birds than to Carnosauria. This chapter examines spinosauroids, carnosaurs, and basal coelurosaurs. Among the taxa in this chapter are the largest carnivorous dinosaurs found in Middle Jurassic through early Late Cretaceous faunal assemblages. The chapter also discusses some of the smallest nonavian dinosaurs, including Compsognathus and Sinosauropteryx.
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The Late Cretaceous tyrannosaurid theropod Alioramus has long been one of the most puzzling large carnivorous dinosaur taxa, largely because for several decades it has been represented only by a single, fragmentary specimen that seems to represent a long-snouted and gracile individual but is difficult to interpret. The discovery of a substantially complete skeleton of Alioramus at the Tsaagan Khuushu locality in the Maastrichtian Nemegt Formation of Mongolia, recovered during the 2001 American Museum–Mongolian Academy of Sciences expedition and described as a new species (Alioramus altai) in 2009, definitively shows that this mysterious taxon is a distinct form of longirostrine tyrannosaurid that lived alongside the larger and more robust Tarbosaurus. Here we describe and figure this remarkably preserved skeleton in detail. We provide exhaustive descriptions and photographs of individual bones, and make extensive comparisons with other tyrannosauroids. This monographic description provides further evidence that Alioramus is an unusual long-snouted, gracile, and slender-limbed taxon with an unpredecented degree of cranial ornamentation among tyrannosaurids and an extremely pneumatized skeleton. Anatomical comparisons indicate that the long skull of Alioramus is an autapomorphic feature that is proportionally longer (relative to femur length) than in any other known tyrannosaurid specimen, including juveniles, and that Alioramus is morphologically distinctive relative to similarly sized individuals of the contemporary and sympatric Tarbosaurus. The holotype specimen of A. altai belongs to a young individual, and many differences between it and the other known specimen of Alioramus (the holotype of A. remotus) may represent ontogenetic variation. The unusual longirostrine skull of Alioramus was largely produced by lengthening of the snout bones (maxilla, nasal, dentary, lacrimal, jugal), rather than the orbiotemporal bones (frontal, postorbital, squamosal, quadratojugal). The long snout, gracile skull bones, comparatively small attachment sites for jaw muscles, and lack of interlocking sutures and a robust orbital brow would have precluded the holotype individual from employing the characteristic “puncture-pull” feeding style of large-bodied adult tyrannosaurids, in which the muscular jaws, thick teeth, and interlocking sutures enabled individuals to bite with enough force to fracture bone. Whether adult Alioramus could utilize “puncture-pull” feeding awaits discovery of mature individuals of the genus. The coexistence of the long-snouted Alioramus and robust and deep-snouted Tarbosaurus, which are found together at the Tsaagan Khuushu locality, demonstrate that multiple large tyrannosaurids were able to live in sympatry, likely because of niche partitioning due to differences in craniofacial morphology and functional behavior.
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The well preserved material of the Late Cretaceous dromaeosaurid, Velociraptor mongoliensis, has allowed us to supplement earlier descriptions of the skull in this species. The skull of V. mongoliensis is similar to that of Deinonychus antirrhopus, but differs from the latter by: (1) laterally convex supratemporal arcade resulting in short, rounded supratemporal fenestra; (2) depressed nasal; (3) longer maxillary process of premaxilla; (4) lack of separate prefrontal, and (5) convex ventral border of the dentary. These differences, especially that in the structure of the temporal region, support generic distinction of Deinonychus and Velociraptor. Skulls of other dromaeosaurids are compared.