A nearly complete skull of the sauropod dinosaur Diamantinasaurus matildae from the Upper Cretaceous Winton Formation of Australia and implications for the early evolution of titanosaurs.

Titanosaurian sauropod dinosaurs were diverse and abundant throughout the Cretaceous, with a global distribution. However, few titanosaurian taxa are represented by multiple skeletons, let alone skulls. Diamantinasaurus matildae, from the lower Upper Cretaceous Winton Formation of Queensland, Australia, was heretofore represented by three specimens, including one that preserves a braincase and several other cranial elements. Herein, we describe a fourth specimen of Diamantinasaurus matildae that preserves a more complete skull-including numerous cranial elements not previously known for this taxon-as well as a partial postcranial skeleton. The skull of Diamantinasaurus matildae shows many similarities to that of the coeval Sarmientosaurus musacchioi from Argentina (e.g. quadratojugal with posterior tongue-like process; braincase with more than one ossified exit for cranial nerve V; compressed-cone-chisel-like teeth), providing further support for the inclusion of both taxa within the clade Diamantinasauria. The replacement teeth within the premaxilla of the new specimen are morphologically congruent with teeth previously attributed to Diamantinasaurus matildae, and Diamantinasauria more broadly, corroborating those referrals. Plesiomorphic characters of the new specimen include a sacrum comprising five vertebrae (also newly demonstrated in the holotype of Diamantinasaurus matildae), rather than the six or more that typify other titanosaurs. However, we demonstrate that there have been a number of independent acquisitions of a six-vertebrae sacrum among Somphospondyli and/or that there have been numerous reversals to a five-vertebrae sacrum, suggesting that sacral count is relatively plastic. Other newly identified plesiomorphic features include: the overall skull shape, which is more similar to brachiosaurids than 'derived' titanosaurs; anterior caudal centra that are amphicoelous, rather than procoelous; and a pedal phalangeal formula estimated as 2-2-3-2-0. These features are consistent with either an early-branching position within Titanosauria, or a position just outside the titanosaurian radiation, for Diamantinasauria, as indicated by alternative character weighting approaches applied in our phylogenetic analyses, and help to shed light on the early assembly of titanosaurian anatomy that has until now been obscured by a poor fossil record.

A new giant sauropod, Australotitan cooperensis gen. et sp. nov., from the mid-Cretaceous of Australia.

A new giant sauropod, Australotitan cooperensis gen. et sp. nov., represents the first record of dinosaurs from the southern-central Winton Formation of the Eromanga Basin, Australia. We estimate the type locality to be 270-300 m from the base of the Winton Formation and compare this to the semi-contemporaneous sauropod taxa, Diamantinasaurus matildae Hocknull et al., 2009, Wintonotitan wattsi Hocknull et al., 2009 and Savannasaurus elliottorum Poropat et al., 2016. The new titanosaurian is the largest dinosaur from Australia as represented by osteological remains and based on limb-size comparisons it reached a size similar to that of the giant titanosaurians from South America. Using 3-D surface scan models we compare features of the appendicular skeleton that differentiate Australotitan cooperensis gen. et sp. nov. as a new taxon. A key limitation to the study of sauropods is the inability to easily and directly compare specimens. Therefore, 3-D cybertypes have become a more standard way to undertake direct comparative assessments. Uncoloured, low resolution, and uncharacterized 3-D surface models can lead to misinterpretations, in particular identification of pre-, syn- and post-depositional distortion. We propose a method for identifying, documenting and illustrating these distortions directly onto the 3-D geometric surface of the models using a colour reference scheme. This new method is repeatable for researchers when observing and documenting specimens including taphonomic alterations and geometric differences. A detailed comparative and preliminary computational phylogenetic assessment supports a shared ancestry for all four Winton Formation taxa, albeit with limited statistical support. Palaeobiogeographical interpretations from these resultant phylogenetic hypotheses remain equivocal due to contrary Asian and South American relationships with the Australian taxa. Temporal and palaeoenvironmental differences between the northern and southern-central sauropod locations are considered to explain the taxonomic and morphological diversity of sauropods from the Winton Formation. Interpretations for this diversity are explored, including an eco-morphocline and/or chronocline across newly developed terrestrial environments as the basin fills.

Myological reconstruction of the pelvic girdle and hind limb of the giant titanosaurian sauropod dinosaur Dreadnoughtus schrani.

Osteological correlates preserve more readily than their soft tissue counterparts in the fossil record; therefore, they can more often provide insight into the soft tissue anatomy of the organism. These insights can in turn elucidate the biology of these extinct organisms. In this study, we reconstruct the pelvic girdle and hind limb musculature of the giant titanosaurian sauropod Dreadnoughtus schrani based on observations of osteological correlates and Extant Phylogenetic Bracket comparisons. Recovered fossils of Dreadnoughtus exhibit remarkably well-preserved, well-developed, and extensive muscle scars. Furthermore, this taxon is significantly larger bodied than any titanosaurian for which a myological reconstruction has previously been performed, rendering this contribution highly informative for the group. All 20 of the muscles investigated in this study are sufficiently well supported to enable reconstruction of at least one division, including reconstruction of the M. ischiocaudalis for the first time in a sauropod dinosaur. In total, 34 osteological correlates were identified on the pelvic girdle and hind limb remains of Dreadnoughtus, allowing the reconstruction of 14 muscles on the basis of Level I or Level II inferences (i.e., not Level I' or Level II' inferences). Comparisons among titanosaurians suggest widespread myological variation, yet potential phylogenetic and other paleobiologic patterns are often obscured by fragmentary preservation, infrequent myological studies, and lack of consensus on the phylogenetic placement of many taxa. However, a ventrolateral accessory process is present on the preacetabular lobe of the ilium in all of the largest titanosauriforms that preserve this skeletal element, suggesting that the presence of this process (representing the origin of the M. puboischiofemoralis internus part II) may be associated with extreme body size. By identifying such myological patterns among titanosauriforms, we can begin to address specific evolutionary and biomechanical questions related to their skeletal anatomy, how they were capable of leaving wide-gauge trackways, and resulting locomotor attributes unique to this clade.

Specialized Craniofacial Anatomy of a Titanosaurian Embryo from Argentina.

The first dinosaur embryos found inside megaloolithid eggs from Auca Mahuevo, Patagonia, were assigned to sauropod dinosaurs that lived approximately 80 million years ago. Discovered some 25 years ago, these considerably flattened specimens still remain the only unquestionable embryonic remains of a sauropod dinosaur providing an initial glimpse into titanosaurian in ovo ontogeny. Here we describe an almost intact embryonic skull, which indicates the early development of stereoscopic vision, and an unusual monocerotic face for a sauropod. The new fossil also reveals a neurovascular sensory system in the premaxilla and a partly calcified braincase, which potentially refines estimates of its prenatal stage. The embryo was found in an egg with thicker eggshell and a partly different geochemical signature than those from the egg-bearing layers described in Auca Mahuevo. The cranial bones are comparably ossified as in previously described specimens but differ in facial anatomy and size. The new specimen reveals significant heterochrony in cranial ossifications when compared with non-sauropod sauropodomorph embryos, and demonstrates that the specialized craniofacial morphology preceded the postnatal transformation of the skull anatomy in adults of related titanosaurians.

Appendicular myological reconstruction of the forelimb of the giant titanosaurian sauropod dinosaur Dreadnoughtus schrani.

Soft tissues are variably preserved in the fossil record with external tissues, such as skin and feathers, more frequently preserved than internal tissues (e.g. muscles). More commonly, soft tissues leave traces of their locations on bones and, for muscles, these clues can be used to reconstruct the musculature of extinct vertebrates, thereby enhancing our understanding of how these organisms moved and the evolution of their locomotor patterns. Herein we reconstruct the forelimb and shoulder girdle musculature of the giant titanosaurian sauropod Dreadnoughtus schrani based on observations of osteological correlates and dissections of taxa comprising the Extant Phylogenetic Bracket of non-avian dinosaurs (crocodilians and birds). Fossils of Dreadnoughtus exhibit remarkably well-preserved, well-developed, and extensive muscle scars. Furthermore, this taxon is significantly larger-bodied than any titanosaurian for which a myological reconstruction has previously been attempted, rendering this myological study highly informative for the clade. In total, 28 muscles were investigated in this study, for which 46 osteological correlates were identified; these osteological correlates allowed the reconstruction of 16 muscles on the basis of Level I or Level II inferences (i.e. not Level I' or Level II' inferences). Comparisons with other titanosaurians suggest widespread myological variation in the clade, although potential phylogenetic patterns are often obscured by fragmentary preservation, infrequent myological studies, and lack of consensus on the systematic position of many taxa. By identifying myological variations within the clade, we can begin to address specific evolutionary and biomechanical questions related to the locomotor evolution in these sauropods.

Osteology and systematics of Uberabatitan ribeiroi (Dinosauria; Sauropoda): a Late Cretaceous titanosaur from Minas Gerais, Brazil.

Uberabatitan ribeiroi is a Late Cretaceous titanosaur (Dinosauria, Sauropoda) from southeastern Brazil. Here we provide a detailed revision of all its available specimens, including new elements from the type-locality. One new autopomorphy is added to diagnosis of the taxon: astragalus with a well-developed anteroposterior crest that mediodistally delimits the tibial articulation. Linear regressions were conducted in an attempt to circumscribe specimens within the type-series, revealing that it is composed of several individuals, with inferred total body lengths varying from 7 to 26 meters. Phylogenetic analyses including U. ribeiroi show that the Brazilian taxon corresponds to a non-saltasaurid lithostrotian titanosaur.

First palaeoneurological study of a sauropod dinosaur from France and its phylogenetic significance.

Despite continuous improvements, our knowledge of the palaeoneurology of sauropod dinosaurs is still deficient. This holds true even for Titanosauria, which is a particularly speciose clade of sauropods with representatives known from numerous Cretaceous sites in many countries on all continents. The data currently available regarding the palaeoneurology of titanosaurs is strongly biased towards Gondwanan forms (Argentina above all, but also India, Malawi and Australia). In contrast, the palaeoneurology of Laurasian titanosaurs is known only from a few taxa from Spain and Uzbekistan, despite the discovery in other countries of Laurasia of a number of neurocranial remains that would lend themselves well to investigations of this kind. To fill in this gap in our knowledge, we subjected a titanosaurian braincase from the uppermost Upper Cretaceous of southern France to X-ray computed tomographic scanning, allowing the generation of 3D renderings of the endocranial cavity enclosing the brain, cranial nerves and blood vessels, as well as the labyrinth of the inner ear. These reconstructions are used to clarify the phylogenetic position of the specimen from the Fox-Amphoux-Métisson site. A combination of characters, including the presence of two hypoglossal rami on the endocast, the average degree of development of the dorsal-head/caudal-middle-cerebral vein system and the relatively short and subequal lengths of the ipsilateral semicircular canals of the labyrinth, are particularly revealing in this respect. They suggest that, compared with the few other Laurasian titanosaurs for which in-depth palaeoneurological data are available, the French taxon is more derived than the distinctly more ancient, possibly non-lithostrotian titanosaur from the Uzbek site of Dzharakuduk but more basal than derived saltasaurids, such as the coeval or slightly more recent forms from the Spanish locality of Lo Hueco.

New information on the Cretaceous sauropod dinosaurs of Zhejiang Province, China: impact on Laurasian titanosauriform phylogeny and biogeography.

Titanosaurs were a globally distributed clade of Cretaceous sauropods. Historically regarded as a primarily Gondwanan radiation, there is a growing number of Eurasian taxa, with several putative titanosaurs contemporaneous with, or even pre-dating, the oldest known Southern Hemisphere remains. The early Late Cretaceous Jinhua Formation, in Zhejiang Province, China, has yielded two putative titanosaurs, Jiangshanosaurus lixianensis and Dongyangosaurus sinensis. Here, we provide a detailed re-description and diagnosis of Jiangshanosaurus, as well as new anatomical information on Dongyangosaurus. Previously, a 'derived' titanosaurian placement for Jiangshanosaurus was primarily based on the presence of procoelous anterior caudal centra. We show that this taxon had amphicoelous anterior-middle caudal centra. Its only titanosaurian synapomorphy is that the dorsal margins of the scapula and coracoid are approximately level with one another. Dongyangosaurus can clearly be differentiated from Jiangshanosaurus, and displays features that indicate a closer relationship to the titanosaur radiation. Revised scores for both taxa are incorporated into an expanded phylogenetic data matrix, comprising 124 taxa scored for 548 characters. Under equal weights parsimony, Jiangshanosaurus is recovered as a member of the non-titanosaurian East Asian somphospondylan clade Euhelopodidae, and Dongyangosaurus lies just outside of Titanosauria. However, when extended implied weighting is applied, both taxa are placed within Titanosauria. Most other 'middle' Cretaceous East Asian sauropods are probably non-titanosaurian somphospondylans, but at least Xianshanosaurus appears to belong to the titanosaur radiation. Our analyses also recover the Early Cretaceous European sauropod Normanniasaurus genceyi as a 'derived' titanosaur, clustering with Gondwanan taxa. These results provide further support for a widespread diversification of titanosaurs by at least the Early Cretaceous.

Late Cretaceous sauropod tooth morphotypes may provide supporting evidence for faunal connections between North Africa and Southern Europe.

The Cretaceous Kem Kem beds of Morocco and equivalent beds in Algeria have produced a rich fossil assemblage, yielding, amongst others, isolated sauropod teeth, which can be used in species diversity studies. These Albian-Cenomanian (∼113-93.9 Ma) strata rarely yield sauropod body fossils, therefore, isolated teeth can help to elucidate the faunal assemblages from North Africa, and their relations with those of contemporaneous beds and geographically close assemblages. Eighteen isolated sauropod teeth from three localities (Erfoud and Taouz, Morocco, and Algeria) are studied here, to assess whether the teeth can be ascribed to a specific clade, and whether different tooth morphotypes can be found in the samples. Two general morphotypes are found, based on enamel wrinkling and general tooth morphology. Morphotype I, with mainly rugose enamel wrinkling, pronounced carinae, lemon-shaped to (sub)cylindrical cross-section and mesiodistal tapering towards an apical tip, shows affinities to titanosauriforms and titanosaurs. Morphotype II, characterized by more smooth enamel, cylindrical cross-section, rectangular teeth with no apical tapering and both labial and lingual wear facets, shows similarities to rebbachisaurids. Moreover, similarities are found between these northwest African tooth morphotypes, and tooth morphotypes from titanosaurs and rebbachisaurids from both contemporaneous finds from north and central Africa, as well as from the latest Cretaceous (Campanian-Maastrichtian, 83.6 Ma-66.0 Ma) of the Ibero-Armorican Island. These results support previous hypotheses from earlier studies on faunal exchange and continental connections between North Africa and Southern Europe in the Cretaceous.

A new giant titanosaur sheds light on body mass evolution among sauropod dinosaurs.

Titanosauria was the most diverse and successful lineage of sauropod dinosaurs. This clade had its major radiation during the middle Early Cretaceous and survived up to the end of that period. Among sauropods, this lineage has the most disparate values of body mass, including the smallest and largest sauropods known. Although recent findings have improved our knowledge on giant titanosaur anatomy, there are still many unknown aspects about their evolution, especially for the most gigantic forms and the evolution of body mass in this clade. Here we describe a new giant titanosaur, which represents the largest species described so far and one of the most complete titanosaurs. Its inclusion in an extended phylogenetic analysis and the optimization of body mass reveals the presence of an endemic clade of giant titanosaurs inhabited Patagonia between the Albian and the Santonian. This clade includes most of the giant species of titanosaurs and represents the major increase in body mass in the history of Titanosauria.

New lower jaw and teeth referred to Maxakalisaurus topai (Titanosauria: Aeolosaurini) and their implications for the phylogeny of titanosaurid sauropods.

Sauropod dinosaurs compose a diversified, well known, and worldwide distributed clade, with a stereotyped body plan: deep trunk, elongated neck and tail, columnar limbs and very small skull. In Brazil, the group is represented by ten formally described Cretaceous species, mostly titanosaurs. This is the case of Maxakalisaurus topai, known based on an incomplete and disarticulated skeleton, unearthed from deposits of the Adamantina Formation in Minas Gerais. Here, we report a partial right dentary, including five isolated teeth, collected from the same site as the type-series of M. topai and tentatively referred to that taxon. The bone is gently curved medially, the functional teeth are set on an anterolingual position, and two replacement teeth are seen per alveoli. New morphological data gathered from that specimen was employed to conduct a comprehensive phylogenetic analysis of Titanosauria (with 42 taxa and 253 characters), based on previous studies. The Aeolosaurini clade was recovered, with Gondwanatitan and Aelosaurus as sister taxa, and Maxakalisaurus, Panamericansaurus, and Rinconsaurus forming a basal polytomy.

Time-calibrated models support congruency between Cretaceous continental rifting and titanosaurian evolutionary history.

Recent model-based phylogenetic approaches have expanded upon the incorporation of extinct lineages and their respective temporal information for calibrating divergence date estimates. Here, model-based methods are explored to estimate divergence dates and ancestral ranges for titanosaurian sauropod dinosaurs, an extinct and globally distributed terrestrial clade that existed during the extensive Cretaceous supercontinental break-up. Our models estimate an Early Cretaceous (approx. 135 Ma) South American origin for Titanosauria. The estimated divergence dates are broadly congruent with Cretaceous geophysical models of supercontinental separation and subsequent continental isolation while obviating the invocation of continuous Late Cretaceous continental connections (e.g. ephemeral land bridges). Divergence dates for mid-Cretaceous African and South American sister lineages support semi-isolated subequatorial African faunas in concordance with the gradual northward separation between South America and Africa. Finally, Late Cretaceous Africa may have linked Laurasian lineages with their sister South American lineages, though the current Late Cretaceous African terrestrial fossil record remains meagre.

Histological evidence for a supraspinous ligament in sauropod dinosaurs.

Supraspinous ossified rods have been reported in the sacra of some derived sauropod dinosaurs. Although different hypotheses have been proposed to explain the origin of this structure, histological evidence has never been provided to support or reject any of them. In order to establish its origin, we analyse and characterize the microstructure of the supraspinous rod of two sauropod dinosaurs from the Upper Cretaceous of Argentina. The supraspinous ossified rod is almost entirely formed by dense Haversian bone. Remains of primary bone consist entirely of an avascular tissue composed of two types of fibre-like structures, which are coarse and longitudinally (parallel to the main axis of the element) oriented. These structures are differentiated on the basis of their optical properties under polarized light. Very thin fibrous strands are also observed in some regions. These small fibres are all oriented parallel to one another but perpendicular to the element main axis. Histological features of the primary bone tissue indicate that the sacral supraspinous rod corresponds to an ossified supraspinous ligament. The formation of this structure appears to have been a non-pathological metaplastic ossification, possibly induced by the continuous tensile forces applied to the element.

Notes on the axial skeleton of the titanosaur Bonitasaura salgadoi (Dinosauria-Sauropoda)

Sauropod axial anatomy is particularly important in understanding morphological features and phylogenetic analyses. Spatial arrangement of zygapophyses and rib articulations, as well as their complex laminar development, help to recognize the relative position of isolated vertebral elements. The presence of anterior, mid and posterior elements along the cervical, dorsal and caudal series in Bonitasaura salgadoi allows the analysis of several anatomical characteristics. These include the pattern of neurocentral closure with unfused, partially fused and completely fused elements in a peculiar temporal sequence, as well as several neural spine modifications and the laminar arrangement. The variations in neural spine morphology include a lateral expansion of the distal tip in cervico-dorsal region, different lateral constituents of these lateral expansions, and a marked deviation in spine angulation. The spinal inclination allows the division into three easily recognizable vertebral regions separated by two landmarks, which adds support in the determination of an accurate vertebral position for isolated elements. Finally, an analysis of the vertebral laminae reveals the importance of examining vertebral series in order to recognize laminar homologies and developmental series. Two vertebral laminae are analyzed here.

A anatomia axial dos saurópodes compõe um conjunto dedados fundamentais para a caracterização morfológica dostáxons, bem como para a determinação de suas relações filogenéticas. A disposição espacial das zigapófises e articulações das costelas, assim como de seu complexo de lâminas acessórias, auxiliam no reconhecimento da posição relativa de elementos vertebrais isolados. A preservação de vértebras anteriores, médias e posteriores nas séries cervical, dorsal e caudal de Bonitasaura salgadoi permite a análise de características adicionais diagnósticas observadas ao longo das séries vertebrais. Estas incluem o grau de fusão da sutura neurocentral,havendo elementos livres, parcialmente fundidos a completamente fundidos em uma sequência temporal peculiar, assim como a presença de diversas modificações do espinho neural e a disposição laminar. As variações na morfologia do espinho neural incluem uma expansão lateral em seu ápice distal na região cervico-dorsal, outros componentes laterais distintos destas expansões, e um desvio crânio-caudal marcado pela angulação do espinho neural. A inclinação espinal permite a divisão em três segmentos vertebrais reconhecíveis separados por duas linhas de transição. Finalmente, a análise das lâminas vertebrais revela a importância de examinar a série vertebral a fim de reconhecer homologias nas lâminas e nas séries de desenvolvimento. Neste trabalho são analisadas duas lâminas vertebrais.

A new titanosaur sauropod from the Atacama Desert, Chile

Partial remains of a titanosaur sauropod collected in the Tolar Formation (Upper Cretaceous) at the Atacama Desert (Antofagasta Region), northern Chile, is described, and a new species, Atacamatitan chilensis gen. et sp. nov., is erected. The material consists mainly of dorsal and caudal vertebrae, part of a humerus and a femur. The presence of a titanosaur confirms the Cretaceous age for the outcrops of red sandstone of the Tolar Formation whose age was previously uncertain, ranging from the Upper Cretaceous to the Paleocene. The new specimen represents the most complete dinosaur reported for this region and one of the most complete titanosaur known from Chile and the pacific margin of South America so far.

Um esqueleto incompleto de um dinossauro titanossaurídeo de depósitos da Formação Tolar (Cretáceo Superior) do deserto do Atacama (região de Antofagasta), norte do Chile é descrito e uma nova espécie, Atacamatitan chilensis gen. et sp. nov., é proposta. O material consiste principalmente de vértebras dorsais e caudais e de partes do úmero e fêmur. A presença de umtitanossauro confirma a idade Cretácea para os afloramentos de arenito avermelhado da Formação Tolar, que eram anteriormente tidos como depositados entre o Cretáceo Superior e o Paleoceno. O exemplar representa o mais completo titanossauro até hoje encontrado na região e um dos mais completos reportados até o momento do Chile e da margem do Oceano Pacífico na América do Sul.

A new Cretaceous terrestrial ecosystem from Gondwana with the description of a new sauropod dinosaur

A unique site at the northern area of Patagonia (Neuquén, Argentina) reveals a terrestrial ecosystem preserved in a detail never reported before in a Late Cretaceous deposit. An extraordinary diversity and abundance of fossils was found concentrated in a 0.5 m horizon in the same quarry, including a new titanosaur sauropod, Futalognkosaurus dukei n.gen., n.sp, which is the most complete giant dinosaur known so far. Several plant leaves, showing a predominance of angiosperms over gymnosperms that likely constituted the diet of F. dukei were found too. Other dinosaurs (sauropods, theropods, ornithopods), crocodylomorphs, pterosaurs, and fishes were also discovered, allowing a partial reconstruction of this Gondwanan continental ecosystem.

Um depósito fóssil na região norte da Patagônia (Neuquén, Argentina) revela um ecossistema nunca antes registrado a este nível de detalhes em depósitos do Cretáceo Superior. Uma diversidade e abundância extraordinária de fósseis encontra-se concentrada em uma camada de 0,5 m no mesmo sítio, incluindo um novo saurópodo titanossaurídeo, Futalognkosaurus dukei n. gen, n. sp., que é o mais completo dinossauro gigante encontrado até a presente data. Foram descobertas váriasfolhas de plantas indicando a predominância de angiospermas sobre gimnospermas que possivelmente formavam a base da dieta de F. dukei. Outros dinossauros (saurópodes, terópodes, ornitópodes), crocodilomorfos, pterossauros e peixes foram também encontrados possibilitando a reconstrução parcialdeste ecossistema continental do Gondwana.