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Concept: Titanosaur


Australian dinosaurs have played a rare but controversial role in the debate surrounding the effect of Gondwanan break-up on Cretaceous dinosaur distribution. Major spatiotemporal gaps in the Gondwanan Cretaceous fossil record, coupled with taxon incompleteness, have hindered research on this effect, especially in Australia. Here we report on two new sauropod specimens from the early Late Cretaceous of Queensland, Australia, that have important implications for Cretaceous dinosaur palaeobiogeography. Savannasaurus elliottorum gen. et sp. nov. comprises one of the most complete Cretaceous sauropod skeletons ever found in Australia, whereas a new specimen of Diamantinasaurus matildae includes the first ever cranial remains of an Australian sauropod. The results of a new phylogenetic analysis, in which both Savannasaurus and Diamantinasaurus are recovered within Titanosauria, were used as the basis for a quantitative palaeobiogeographical analysis of macronarian sauropods. Titanosaurs achieved a worldwide distribution by at least 125 million years ago, suggesting that mid-Cretaceous Australian sauropods represent remnants of clades which were widespread during the Early Cretaceous. These lineages would have entered Australasia via dispersal from South America, presumably across Antarctica. High latitude sauropod dispersal might have been facilitated by Albian-Turonian warming that lifted a palaeoclimatic dispersal barrier between Antarctica and South America.

Concepts: Continent, Pacific Ocean, Cretaceous, Antarctica, Sauropoda, Dinosaur, Titanosaur, Macronaria


Sauropod dinosaurs are a group of herbivorous dinosaurs which exceeded all other terrestrial vertebrates in mean and maximal body size. Sauropod dinosaurs were also the most successful and long-lived herbivorous tetrapod clade, but no abiological factors such as global environmental parameters conducive to their gigantism can be identified. These facts justify major efforts by evolutionary biologists and paleontologists to understand sauropods as living animals and to explain their evolutionary success and uniquely gigantic body size. Contributions to this research program have come from many fields and can be synthesized into a biological evolutionary cascade model of sauropod dinosaur gigantism (sauropod gigantism ECM). This review focuses on the sauropod gigantism ECM, providing an updated version based on the contributions to the PLoS ONE sauropod gigantism collection and on other very recent published evidence. The model consist of five separate evolutionary cascades (“Reproduction”, “Feeding”, “Head and neck”, “Avian-style lung”, and “Metabolism”). Each cascade starts with observed or inferred basal traits that either may be plesiomorphic or derived at the level of Sauropoda. Each trait confers hypothetical selective advantages which permit the evolution of the next trait. Feedback loops in the ECM consist of selective advantages originating from traits higher in the cascades but affecting lower traits. All cascades end in the trait “Very high body mass”. Each cascade is linked to at least one other cascade. Important plesiomorphic traits of sauropod dinosaurs that entered the model were ovipary as well as no mastication of food. Important evolutionary innovations (derived traits) were an avian-style respiratory system and an elevated basal metabolic rate. Comparison with other tetrapod lineages identifies factors limiting body size.

Concepts: Evolution, Metabolism, Bird, Paleontology, Sauropoda, Dinosaur, Sauropodomorpha, Titanosaur


We describe Sarmientosaurus musacchioi gen. et sp. nov., a titanosaurian sauropod dinosaur from the Upper Cretaceous (Cenomanian-Turonian) Lower Member of the Bajo Barreal Formation of southern Chubut Province in central Patagonia, Argentina. The holotypic and only known specimen consists of an articulated, virtually complete skull and part of the cranial and middle cervical series. Sarmientosaurus exhibits the following distinctive features that we interpret as autapomorphies: (1) maximum diameter of orbit nearly 40% rostrocaudal length of cranium; (2) complex maxilla-lacrimal articulation, in which the lacrimal clasps the ascending ramus of the maxilla; (3) medial edge of caudal sector of maxillary ascending ramus bordering bony nasal aperture with low but distinct ridge; (4) ‘tongue-like’ ventral process of quadratojugal that overlaps quadrate caudally; (5) separate foramina for all three branches of the trigeminal nerve; (6) absence of median venous canal connecting infundibular region to ventral part of brainstem; (7) subvertical premaxillary, procumbent maxillary, and recumbent dentary teeth; (8) cervical vertebrae with ‘strut-like’ centroprezygapophyseal laminae; (9) extremely elongate and slender ossified tendon positioned ventrolateral to cervical vertebrae and ribs. The cranial endocast of Sarmientosaurus preserves some of the most complete information obtained to date regarding the brain and sensory systems of sauropods. Phylogenetic analysis recovers the new taxon as a basal member of Lithostrotia, as the most plesiomorphic titanosaurian to be preserved with a complete skull. Sarmientosaurus provides a wealth of new cranial evidence that reaffirms the close relationship of titanosaurs to Brachiosauridae. Moreover, the presence of the relatively derived lithostrotian Tapuiasaurus in Aptian deposits indicates that the new Patagonian genus represents a ‘ghost lineage’ with a comparatively plesiomorphic craniodental form, the evolutionary history of which is missing for at least 13 million years of the Cretaceous. The skull anatomy of Sarmientosaurus suggests that multiple titanosaurian species with dissimilar cranial structures coexisted in the early Late Cretaceous of southern South America. Furthermore, the new taxon possesses a number of distinctive morphologies-such as the ossified cervical tendon, extremely pneumatized cervical vertebrae, and a habitually downward-facing snout-that have rarely, if ever, been documented in other titanosaurs, thus broadening our understanding of the anatomical diversity of this remarkable sauropod clade. The latter two features were convergently acquired by at least one penecontemporaneous diplodocoid, and may represent mutual specializations for consuming low-growing vegetation.

Concepts: Skull, Cranial nerves, Cretaceous, Patagonia, Sauropoda, Dinosaur, Titanosaur, Maxillary nerve


Titanosaurian sauropod dinosaurs were the most diverse and abundant large-bodied herbivores in the southern continents during the final 30 million years of the Mesozoic Era. Several titanosaur species are regarded as the most massive land-living animals yet discovered; nevertheless, nearly all of these giant titanosaurs are known only from very incomplete fossils, hindering a detailed understanding of their anatomy. Here we describe a new and gigantic titanosaur, Dreadnoughtus schrani, from Upper Cretaceous sediments in southern Patagonia, Argentina. Represented by approximately 70% of the postcranial skeleton, plus craniodental remains, Dreadnoughtus is the most complete giant titanosaur yet discovered, and provides new insight into the morphology and evolutionary history of these colossal animals. Furthermore, despite its estimated mass of about 59.3 metric tons, the bone histology of the Dreadnoughtus type specimen reveals that this individual was still growing at the time of death.

Concepts: Biology, Reptile, Mammal, Cretaceous, Sauropoda, Dinosaur, Titanosaur, Plesiosaur


Titanosauria is an exceptionally diverse, globally-distributed clade of sauropod dinosaurs that includes the largest known land animals. Knowledge of titanosaurian pedal structure is critical to understanding the stance and locomotion of these enormous herbivores and, by extension, gigantic terrestrial vertebrates as a whole. However, completely preserved pedes are extremely rare among Titanosauria, especially as regards the truly giant members of the group. Here we describe Notocolossus gonzalezparejasi gen. et sp. nov. from the Upper Cretaceous of Mendoza Province, Argentina. With a powerfully-constructed humerus 1.76 m in length, Notocolossus is one of the largest known dinosaurs. Furthermore, the complete pes of the new taxon exhibits a strikingly compact, homogeneous metatarsus-seemingly adapted for bearing extraordinary weight-and truncated unguals, morphologies that are otherwise unknown in Sauropoda. The pes underwent a near-progressive reduction in the number of phalanges along the line to derived titanosaurs, eventually resulting in the reduced hind foot of these sauropods.

Concepts: Sauropoda, Dinosaur, Sauropodomorpha, Diplodocus, Saurischia, Titanosaur, Bruhathkayosaurus, Macronaria


Increased excavation of dinosaurs from China over the last two decades has enriched the record of Asian titanosauriform sauropods. However, the relationships of these sauropods remain contentious, and hinges on a few well-preserved taxa, such as Euhelopus zdanskyi. Here we describe a new sauropod, Yongjinglong datangi gen. nov. et sp. nov., from the Lower Cretaceous Hekou Group in the Lanzhou Basin of Gansu Province, northwestern China. Yongjinglong datangi is characterized by the following unique combination of characters, including seven autapomorphies: long-crowned, spoon-shaped premaxillary tooth; axially elongate parapophyses on the cervical vertebra; very deep lateral pneumatic foramina on the lateral surfaces of the cervical and cranial dorsal vertebral centra; low, unbifurcated neural spine fused with the postzygapophyses to form a cranially-pointing, triangular plate in a middle dorsal vertebra; an “XI”-shaped configuration of the laminae on the arches of the middle dorsal vertebrae; a very long scapular blade with straight cranial and caudal edges; and a tall, deep groove on the lateral surface of the distal shaft of the radius. The new specimen shares several features with other sauropods: a pronounced M. triceps longus tubercle on the scapula and ventrolaterally elongated parapophyses in its cervical vertebra as in Euhelopodidae. Based on phylogenetic analyses Yongjinglong datangi is highly derived within Titanosauria, which suggests either a remarkable convergence with more basal titanosauriform sauropods in the Early Cretaceous or a retention of plesiomorphic features that were lost in other titanosaurians. The morphology and remarkable length of the scapulocoracoid reveal an unusual relationship between the shoulder and the middle trunk: the scapulocoracoid spans over half of the length of the trunk. The medial, notch-shaped coracoid foramen and the partially fused scapulocoracoid synostosis suggest that the specimen is a subadult individual. This specimen sheds new light on the diversity of Early Cretaceous Titanosauriformes in China.

Concepts: Vertebral column, Vertebra, Cretaceous, Cervical vertebrae, Sauropoda, Dinosaur, Titanosaur, Euhelopus


Sauropod dinosaurs exhibit the largest ontogenetic size range among terrestrial vertebrates, but a dearth of very young individuals has hindered understanding of the beginning of their growth trajectory. A new specimen of Rapetosaurus krausei sheds light on early life in the smallest stage of one of the largest dinosaurs. Bones record rapid growth rates and hatching lines, indicating that this individual weighed ~3.4 kilograms at hatching. Just several weeks later, when it likely succumbed to starvation in a drought-stressed ecosystem, it had reached a mass of ~40 kilograms and was ~35 centimeters tall at the hip. Unexpectedly, Rapetosaurus limb bones grew isometrically throughout their development. Cortical remodeling, limb isometry, and thin calcified hypertrophic metaphyseal cartilages indicate an active, precocial growth strategy.

Concepts: Developmental biology, Bird, Sauropoda, Dinosaur, Titanosaur, Tyrannosaurus, Argentinosaurus, Rapetosaurus


The sauropod of El Oterillo II is a specimen that was excavated from the Castrillo de la Reina Formation (Burgos, Spain), late Barremian-early Aptian, in the 2000s but initially remained undescribed. A tooth and elements of the axial skeleton, and the scapular and pelvic girdle, represent it. It is one of the most complete titanosauriform sauropods from the Early Cretaceous of Europe and presents an opportunity to deepen our understanding of the radiation of this clade in the Early Cretaceous and study the paleobiogeographical relationships of Iberia with Gondwana and with other parts of Laurasia. The late Barremian-early Aptian is the time interval in the Cretaceous with the greatest diversity of sauropod taxa described in Iberia: two titanosauriforms, Tastavinsaurus and Europatitan; and a rebbachisaurid, Demandasaurus. The new sauropod Europatitan eastwoodi n. gen. n. sp. presents a series of autapomorphic characters in the presacral vertebrae and scapula that distinguish it from the other sauropods of the Early Cretaceous of Iberia. Our phylogenetic study locates Europatitan as the basalmost member of the Somphospondyli, clearly differentiated from other clades such as Brachiosauridae and Titanosauria, and distantly related to the contemporaneous Tastavinsaurus. Europatitan could be a representative of a Eurogondwanan fauna like Demandasaurus, the other sauropod described from the Castrillo de la Reina Formation. The presence of a sauropod fauna with marked Gondwananan affinities in the Aptian of Iberia reinforces the idea of faunal exchanges between this continental masses during the Early Cretaceous. Further specimens and more detailed analysis are needed to elucidate if this Aptian fauna is caused by the presence of previously unnoticed Aptian land bridges, or it represents a relict fauna from an earlier dispersal event.

Concepts: Phylogenetic nomenclature, Cladistics, Cretaceous, Sauropoda, Dinosaur, Titanosaur, Psittacosaurus, Early Cretaceous


Dermal armor is one of the most intriguing features of some titanosaurs, the only sauropod dinosaurs that bore osteoderms. Some studies have revealed cavities of varying sizes inside some titanosaur osteoderms, interpreted as the result of bone remodeling for mineral mobilization. Several hypotheses have been proposed to explain the need for mineral mobilization in titanosaurs. However, rejecting those hypotheses was difficult with hitherto available evidence. The Upper Cretaceous site of Lo Hueco (Cuenca; Spain) has yielded one of the largest titanosaur osteoderm sets available. Observation of pre-existing breakages in the fossils and CT-scanning have revealed a predominant internal channel network for bulb and root osteoderms: most had a very compact spongy bone core, perfused by large longitudinal branching neurovascular canals. Only few osteoderms from the same bed, which was deposited in a single and short event, had areas with low-density spongy bone. This void-like low-density bone is always associated with internal channels. It is also present in osteoderms of different sizes. This scenario is best explained when considering that Lo Hueco titanosaurs might have used their osteoderms as a source of calcium that was mobilized during oogenesis, although other hypotheses cannot be completely ruled out.

Concepts: Scientific method, Bone, Skeletal system, Theory, Sauropoda, Dinosaur, Titanosaur, Saltasaurus


Titanosaurians were a flourishing group of sauropod dinosaurs during Cretaceous times. Fossils of titanosaurians have been found on all continents and their remains are abundant in a number of Late Cretaceous sites. Nonetheless, the cranial anatomy of titanosaurians is still very poorly known. The Spanish latest Cretaceous locality of “Lo Hueco” yielded a relatively well preserved, titanosaurian braincase, which shares a number of phylogenetically restricted characters with Ampelosaurus atacis from France such as a flat occipital region. However, it appears to differ from A. atacis in some traits such as the greater degree of dorsoventral compression and the presence of proatlas facets. The specimen is, therefore, provisionally identified as Ampelosaurus sp. It was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. Our investigation highlights that, although titanosaurs were derived sauropods with a successful evolutionary history, they present a remarkably modest level of paleoneurological organization. Compared with the condition in the basal titanosauriform Giraffatitan brancai, the labyrinth of Ampelosaurus sp. shows a reduced morphology. The latter feature is possibly related to a restricted range of head-turning movements.

Concepts: Sauropoda, Dinosaur, Titanosaur