Cranial bones and atlas of titanosaurs (Dinosauria,Sauropoda) from Late Cretaceous (Bauru Group) of Uberaba,Minas Gerais State,Brazil

Isolated left prefrontal, left squamosal and atlas of titanosaur dinosaurs are described and compared. They come from the Late Cretaceous Serra da Galga Member of the Marília Formation at the Serra do Veadinho region, Peirópolis (Uberaba County, Minas Gerais State, Brazil). Due to the sparse cranial elements of titanosaurs already known from Brazil, these specimens are noticeable to be presented. In addition, the atlas vertebra is described for the first time for Brazilian titanosaurs. The morphology of the cranial bones closely resembles lithostratian titanosaurs, such as Rapetosaurus, rather than basal titanosaurs. The atlas is similar to that of other titanosaurs, suggesting that the anatomy of this element seems to be more conservative than other vertebral elements, in which vertebral laminae play an important rule in titanosaur taxonomy.     

A new titanosaur from the Lower Cretaceous of Brazil

Here we describe a new record of a sauropod dinosaur from the Lower Cretaceous (Hauterivian–Barremian) Rio Piranhas Formation, Sousa Basin, NE Brazil. Dinosaur fossil bones from this deposit were unknown until now. Thus, the discovery of a sauropod fibula from this locality is highly significant. Our discovery represents an indeterminate titanosaur and the earliest stratigraphic occurrence of this group in central Gondwana. When compared to chronocorrelate titanosaur trackmakers of this geological unit, this fossil specimen appears substantially smaller. Histological analysis of the fibula suggests that this is a relatively young individual (approximately 40–50% adult body size) that had passed its most rapid phase of early juvenile growth, but had not yet attained somatic maturity. Thus, the fibula recovered is from a young individual rather than from a small-bodied adult titanosaur.     

New cranial remains of titanosaurian sauropod dinosaurs from the Late Cretaceous of Fox-Amphoux-Métisson (Var,SE France)

The Fox-Amphoux Syncline of the Aix Basin (SE France) has yielded a diverse Late Cretaceous vertebrate assemblage, including several taxa of dinosaurs. Here, we report on cranial material of titanosaurian sauropods, which consist of a partial braincase, two fragmentary skull roofs, and nine teeth, from the Métisson locality (Var Department). The braincase differs from those found previously in Europe (i.e., Ampelosaurus atacis, Lirainosaurus astibiae, and an unnamed juvenile skull from Romania) as well as from other titanosaurian braincases in having a groove that extends along the ventral surface of the occipital condyle neck (this feature may be autapomorphic). One of the fragmentary skull roofs may belong to the same taxon, whereas the other suggests the presence of a second titanosaur at Métisson very close to Ampelosaurus. Two dental morphologies are present in the sample; one of them includes teeth of different sizes. We suggest that this could be accounted for by age differences within a single taxon, or be due to different positions in the tooth row. The presence of a new, still unnamed titanosaurian taxon in the Ibero-Armorican Island supports previous works indicating a high titanosaurian diversity during the Campanian-Maastrichtian in southern Europe.     

Multiple neoplasms in a single sauropod dinosaur from the Upper Cretaceous of Brazil

Tumors are thought rare in dinosaurs, previously limited in distribution to a single family – hadrosaurs. Here we recognized two different neoplastic lesions, osteoma and hemangioma, in a single caudal vertebra of a titanosaur dinosaur from the Upper Cretaceous of Brazil, diagnosed on the basis of macroscopic, radiological and histological analysis. This is the first occurrence of neoplasia in a non-hadrosaur dinosaur. Therefore, neoplasia is not restricted to hadrosaur dinosaurs, as previously suggested.     

Multiple paleopathologies in the dinosaur Bonitasaura salgadoi (Sauropoda: Titanosauria) from the Upper Cretaceous of Patagonia,Argentina

Bones in the fossil record sometimes exhibit unusual structures that can be attributed to pathologies, taphonomic alterations, or morphological variation. The goal of this work is to describe three bone abnormalities present in the type specimen of Bonitasaura salgadoi from the Cretaceous of north Patagonia, Argentina. The studied material corresponds to a left femur, a left metatarsal III, and a right prezygapophysis of a mid-caudal vertebra. Macroscopic and/or histological examinations were conducted, and a discussion concerning the origin of each abnormality is provided. The results of this study suggest that the anomalous structures are pathologies. In the femur, an osteoblastic tumor was identified by the presence of a large outgrowth of ovoid appearance with a spiculated microstructural pattern. The metatarsal III shows an enthesophyte (or bone spur) based on its location, shape and growth in parallel to the long axis of the bone element. Finally, the abnormal tissue observed in the prezygapophysis of the caudal vertebra was determined to be an infection by the presence of reactive new bone associated with a local widening of the subperiosteal margin and a sinus of drainage. This is the first report of multiple pathologies in a single specimen of a titanosaur, and it provides new insights about paleopathologies in sauropod dinosaurs.     

Review of Cretaceous sauropod dinosaurs from Central Asia

There are 24 known localities for skeletal remains of sauropod dinosaurs in the republics of Central Asia (Kazakhstan, Kyrgyzstan, Tajikistan, and Uzbekistan). Sauropod remains are very rare at all these localities and represented usually only by isolated teeth. Only narrow-crowned teeth are known from the Cretaceous of Central Asia. The oldest record of such teeth is from the Aptian Sultanbobo Formation of Uzbekistan. Exposures of the Turonian Bissekty Formation at the most productive vertebrate locality in the region, Dzharakuduk in Uzbekistan, has yielded many isolated teeth and a few skeletal remains that can be attributed to a non-lithostrotian titanosaur. Similar narrow-crowned, cylindrical teeth from Cenomanian-to Coniacian-age strata in the Kyzylkum Desert, Uzbekistan, may belong to a closely related taxon. Another taxon, with teeth that are pentagonal in cross-section, is known from the Santonian Yalovach and Bostobe formations of Tajikistan and western Kazakhstan, respectively. A femur reported from the Santonian Syuksyuk Formation of southern Kazakhstan possibly belongs to a lithostrotian titanosaur. The change in tooth structure at the Coniacian–Santonian boundary in the region possibly suggests replacement of non-lithostrotian titanosaurs by lithostrotians. The titanosaur from the Bissekty Formation is similar to Dongyangosaurus sinensis from the Cenomanian–Turonian of Zhejiang (China) in the extensive pneumatization of the neural arch on the anterior caudal vertebrae with several fossae. It also resembles Baotianmansaurus henanensis from the Cenomanian of Henan (China) in the possession of very short anterior caudal centra. These three taxa possibly represent an as yet formally unrecognized endemic clade of Asian non-lithostrotian titanosaurs.     

A stegosaur tooth (Dinosauria, Ornithischia) from the Early Cretaceous of southwestern France

The spotty nature of the terrestrial fossil record for the Mesozoic hinders a more complete understanding of dinosaur diversity. For stegosaurs (Ornithischia), the plated dinosaurs, only a few and fragmentary remains are reported from the Early Cretaceous of Europe. A recent revision concluded that only a partial vertebra of the nomen dubium Craterosaurus (?Aptian, England) could be considered as stegosaurian. Here we report on a stegosaur tooth from the Early Cretaceous (Berriasian) Purbeckian deposits of Cherves-de-Cognac (Charente), southwestern France. This tiny tooth was examined in detail using microtomography. Comparisons being limited by the rarity of stegosaur tooth rows material (e.g., from the skull of the holotype of Stegosaurus stenops) and dental material, notably from Europe, we observed new material of cf. Stegosaurus armatus and Hesperosaurus mjosi from the Upper Jurassic Morrison Formation of Wyoming (USA). The tooth shows the most similarities to the Late Jurassic genera Stegosaurus and Hesperosaurus, but differs in having a distinctive downwardly arched (V-shaped) cingulum on the ?lingual face (maxillary tooth hypothesis). It is referred to as Stegosauria indeterminate, a medium-sized quadrupedal herbivore that inhabited an emerged land between the Armorican Massif and the Massif Central. This finding is the first evidence of a stegosaur from the Early Cretaceous of France and a welcome addition to the meagre European record of that time. In addition, it is the second stegosaurian tooth crown reported from Europe. The assemblage of ornithischians of Cherves-de-Cognac shares some similarities with that of the Early Cretaceous (Berriasian) of the Purbeck Limestone Group, southern England. The relative rarity of ornithischian osteological remains in both Purbeckian environments suggests that most of these dinosaurs were mainly inhabitants of inland terrestrial palaeoenvironments.     

Additions to the sauropod dinosaur fauna of the Cenomanian (early Late Cretaceous) Kem Kem beds of Morocco: Palaeobiogeographical implications of the mid-Cretaceous African sauropod fossil record

Determining the relationships of mid-Late Cretaceous African taxa is central to understanding the timing and resultant palaeobiogeographical patterns of Gondwanan fragmentation. The early Late Cretaceous (Cenomanian) Kem Kem beds of southeastern Morocco preserve a diverse vertebrate fauna, including sauropod dinosaurs. Sauropod material includes the holotype of the rebbachisaurid diplodocoid Rebbachisaurus garasbae and fragmentary remains representing indeterminate titanosauriforms and rebbachisaurids. Here, we describe two new specimens from the Kem Kem beds. A dorsal neural arch with complex internal pneumaticity is tentatively attributed to a somphospondylan titanosauriform. A caudal vertebra possessing several rebbachisaurid synapomorphies is excavated by a large and pervasive lateral pneumatic foramen, a feature undocumented in other rebbachisaurids. However, caudal vertebrae are currently unknown for the sympatric R. garasbae, so this element could be referable to that taxon or a second, previously unknown, rebbachisaurid species. Interestingly, this new caudal vertebra displays a mosaic of features otherwise restricted to limaysaurine or nigersaurine rebbachisaurids, suggesting a placement basal to these clades, which is the position usually recovered for R. garasbae in phylogenetic analyses. A review of the mid-Cretaceous African sauropod fossil record removes the Cretaceous record of dicraeosaurids from Africa, restricting this clade to a single post-Jurassic occurrence in Argentina. All diagnostic sauropod remains can be attributed to titanosauriforms or rebbachisaurids. Whereas rebbachisaurids were seemingly restricted to northwestern Africa and disappeared post-Cenomanian, titanosauriforms were widespread across the African continent and survived until the latest Cretaceous. The development of the mid-Cretaceous Trans-Saharan Seaway might have acted as a dispersal barrier for rebbachisaurids and other vertebrate groups. In contrast, titanosauriforms might have been able to cross this barrier, but it is possible that they were also unable to disperse, and that northwestern African titanosauriforms were not closely related to taxa from the rest of the African continent. New materials and a better understanding of titanosaur interrelationships will be crucial in teasing these scenarios apart.     

A complete titanosaur femur from West Texas with comments regarding hindlimb posture

An isolated titanosaur femur recovered from the Javelina Formation (Maastrichtian) of Big Bend National Park, Texas is the most complete example yet reported from North America. The specimen is likely referable to Alamosaurus sanjuanensis, the only titanosaur thus far known from Upper Cretaceous strata in North America, but cannot be attributed with certainty to that taxon. Compared to femora from other titanosaurs, the specimen has a relatively reduced abductor crest, a less elevated femoral head, and a distal joint surface that is orthogonal to the long axis of the shaft. These differences suggest that the Big Bend femur pertains to a species where hindlimb stance was closer to vertical, and with a comparatively narrower gait than other titanosaurids.     

Preservation of large titanosaur sauropods in overbank fluvial facies: A case study in the Cretaceous of Argentina

Patagonia exhibits a particularly abundant record of Cretaceous dinosaurs with worldwide relevance. Although paleontological studies are relatively numerous, few include taphonomic information about these faunas. This contribution provides the first detailed sedimentological and taphonomical analyses of a dinosaur bone quarry from northern Neuquén Basin. At Arroyo Seco (Mendoza Province, Argentina), a large parautochthonous/autochthonous accumulation of articulated and disarticulated bones that represent several sauropod individuals has been discovered. The fossil remains, assigned to Mendozasaurus neguyelap González Riga, correspond to a large (18–27-m long) sauropod titanosaur collected in the strata of the Río Neuquén Subgroup (late Turoronian–late Coniacian). A taphonomic viewpoint recognizes a two-fold division into biostratinomic and fossil-diagenetic processes. Biostratinomic processes include (1) subaerial biodegradation of sauropod carcasses on well-drained floodplains, (2) partial or total skeletal disarticulation, (3) reorientation of bones by sporadic overbank flows, and (4) subaerial weathering. Fossil-diagenetic processes include (1) plastic deformation of bones, (2) initial permineralization with hematite, (3) fracturing and brittle deformation due to lithostatic pressure; (4) secondary permineralization with calcite in vascular canals and fractures, and (5) postfossilization bone weathering. This type of bone concentration, also present in Rincón de los Sauces (northern Patagonia), suggests that overbank facies tended to accumulate large titanosaur bones. This taphonomic mode, referred to as “overbank bone assemblages”, outlines the potential of crevasse splay facies as important sources of paleontological data in Cretaceous meandering fluvial systems.     

A titanosaur (Dinosauria: Sauropoda) osteoderm from the Alcântara Formation (Cenomanian), São Luís Basin,Northeastern Brazil

Among titanosaurs, osteoderms are morphologically diverse and have been reported from deposits in South America, Africa, Madagascar, Eurasia and Oceania. In Brazil, titanosaur osteoderms are rare and have only been recorded from Bauru Basin sedimentary rocks. Here, we describe a keeled titanosaur osteoderm, which is the first occurrence of an osteoderm specimen in the São Luís Basin, Northeastern Brazil. This osteoderm is characterized by an external bulb, an internal root, and a very rugose cingulum limiting the external and internal surfaces. These characteristics are typical of Titanosauria dermal bones, and this specimen strengthens the idea that the armored sauropods were present during the early Late Cretaceous of Northeastern Brazil, extending their distribution in the northernmost portion of South America.     

Fossils Explained 83: Diversity of skull morphology in the Titanosauria

Titanosaurs were a globally distributed group of sauropod dinosaurs. They had diverse forms and a wide-gauge stance, with a few of their species reaching immense sizes, such as Argentinosaurus huinculensis and Patagotitan mayorum (reaching >35 m in length). There are about 100 valid titanosaur species known so far, but most of the originally described species are no longer valid, due to the incomplete nature of fossil materials. Our understanding of titanosaur skull morphology is based on very few incomplete fragmented cranial materials and findings of the complete skull are even rarer. Understanding the skull morphology of extinct animals helps palaeontologists make deductions of feeding mechanisms and also provide an idea about their appearance when they were alive. Diversity in titanosaur skull morphology is greater than that of any other sauropod clade, indicating diversity in feeding mechanism among these dinosaurs. Titanosaurs were the last surviving clade of sauropod dinosaurs, occupying nearly every ecological niche around the world during the Late Cretaceous, and resulting in a rich diversity in this group. This article highlights diversity in the basic structure of sauropods with special emphasis on titanosaur skull morphology.     

Titanosaur (Dinosauria: Sauropoda) Osteoderms from the Maastrichtian of Uberaba, Minas Gerais State, Brazil

Titanosaur remains are common findings at the paleontological site of Peirópolis in Uberaba. Among those remains, two osteoderms referred to titanosaur sauropods were reported. Both dermal bones share many features, such as the coarse texture, parallel ventral grooves, lack of the cingulum and presence of a ventral ridge (crest). Armored titanosaurs had wide distribution in the Upper Cretaceous, and have been reported from Argentina, Brazil, France, Madagascar, Malawi, Romania and Spain. In the present work we describe and compare the Brazilian titanosaur osteoderms found thus far.     

The carbonate fraction in carbonatitic Italian lamprophyres

Alkaline and ultramafic lamprophyres represent the majority of pre-Pleistocene alkaline mafic magmatic activity in Italy and have been described from several localities. The age of magmatism ranges from Triassic to Lower Oligocene. Some contain appreciable amounts of carbonate. The primary carbonate of the Italian carbonatitic lamprophyres is mainly Sr- or Mn-rich calcite that occurs mostly as immiscible ocelli or as groundmass. Its textural occurrence, composition, and relationship with co-precipitating silicate phases is taken as evidence of an igneous origin. Low BaO and REE contents in the carbonate are explained by early crystallization of essential mica and subordinate apatite. Whole rock analyses and isotopic data (Rukhlov, A.S., Bell, K., Vichi, G., Stoppa, F., submitted for publication. The heterogeneous deep mantle: the Sr, Pb and Nd isotopic evidence from Early Cretaceous alkaline lamprophyres of Southern Tuscany, Italy. Lithos.) suggest a mantle origin for these rocks and rule out contamination in either high or low pressure regimes. The bulk compositions of the carbonatitic lamprophyres have high HFSE / LILE and LREE / HREE ratios and although the abundances of these elements are generally lower than for carbonatites s.s., they are comparable with the abundances in other ‘carbonate-free’ Italian lamprophyres and Italian carbonatites, suggesting similar mantle sources. Moreover, the age of the Italian lamprophyres, ranging from Middle Triassic to Lower Oligocene, is much greater than the Pleistocene age of Italian carbonatites and indicates that the source remained similar over a long time span.     

Titanosaur (Dinosauria: Sauropoda) Osteoderms from the Maastrichtian of Uberaba,Minas Gerais State,Brazil

Titanosaur remains are common findings at the paleontological site of Peirópolis in Uberaba. Among those remains, two osteoderms referred to titanosaur sauropods were reported. Both dermal bones share many features, such as the coarse texture, parallel ventral grooves, lack of the cingulum and presence of a ventral ridge (crest). Armored titanosaurs had wide distribution in the Upper Cretaceous, and have been reported from Argentina, Brazil, France, Madagascar, Malawi, Romania and Spain. In the present work we describe and compare the Brazilian titanosaur osteoderms found thus far.     

New rebbachisaurid (Dinosauria: Sauropoda) material from the Wessex Formation (Barremian, Early Cretaceous), Isle of Wight, United Kingdom

Rebbachisauridae is a poorly understood clade of diplodocoid sauropod dinosaurs, currently known only from the Cretaceous of Africa, Europe and South America. European representatives are particularly rare and fragmentary. Here, we report an anterior caudal vertebra from the Barremian (Early Cretaceous) Wessex Formation of the Isle of Wight, off the southern coast of England. This specimen possesses several features known only in rebbachisaurids and shares two synapomorphies with the Afro-European taxa Demandasaurus darwini and Nigersaurus taqueti, both pertaining to the morphology of the neural spine. These features are the development of triangular lateral processes and the presence of an elliptical fossa on the lateral surface, bounded by the lateral lamina and postspinal rugosity. The Isle of Wight specimen also shares several features solely with Demandasaurus, indicating a close relationship with the Spanish taxon. These include the presence of a hyposphenal ridge, as well as an anteriorly excavated caudal rib that is restricted almost entirely to the neural arch. However, it differs from Demandasaurus in a number of ways, including the lack of excavation on the posterior surface of the caudal rib, the orientation of the neural spine, and the composition and morphology of the lateral lamina. In addition, the Isle of Wight vertebra possesses one potential autapomorphy: bifurcation of the elliptical fossa on the neural spine. However, because of the fragmentary nature of the material, a new name is not erected. Along with Demandasaurus and Histriasaurus boscarollii, this caudal vertebra indicates the presence of at least three European rebbachisaurid taxa and provides new anatomical information on this enigmatic clade of sauropod dinosaurs.     

New rebbachisaurid (Dinosauria: Sauropoda) material from the Wessex Formation (Barremian,Early Cretaceous), Isle of Wight,United Kingdom

Rebbachisauridae is a poorly understood clade of diplodocoid sauropod dinosaurs, currently known only from the Cretaceous of Africa, Europe and South America. European representatives are particularly rare and fragmentary. Here, we report an anterior caudal vertebra from the Barremian (Early Cretaceous) Wessex Formation of the Isle of Wight, off the southern coast of England. This specimen possesses several features known only in rebbachisaurids and shares two synapomorphies with the Afro-European taxa Demandasaurus darwini and Nigersaurus taqueti, both pertaining to the morphology of the neural spine. These features are the development of triangular lateral processes and the presence of an elliptical fossa on the lateral surface, bounded by the lateral lamina and postspinal rugosity. The Isle of Wight specimen also shares several features solely with Demandasaurus, indicating a close relationship with the Spanish taxon. These include the presence of a hyposphenal ridge, as well as an anteriorly excavated caudal rib that is restricted almost entirely to the neural arch. However, it differs from Demandasaurus in a number of ways, including the lack of excavation on the posterior surface of the caudal rib, the orientation of the neural spine, and the composition and morphology of the lateral lamina. In addition, the Isle of Wight vertebra possesses one potential autapomorphy: bifurcation of the elliptical fossa on the neural spine. However, because of the fragmentary nature of the material, a new name is not erected. Along with Demandasaurus and Histriasaurus boscarollii, this caudal vertebra indicates the presence of at least three European rebbachisaurid taxa and provides new anatomical information on this enigmatic clade of sauropod dinosaurs.     

Tetrapods from the Upper Cretaceous (Turonian–Maastrichtian) Bauru Group of Brazil: a reappraisal

An updated, annotated list of all tetrapods from the Adamantina, Uberaba and Marília formations (Bauru Group), which constitute some of the best studied Upper Cretaceous units in Brazil, is presented. Tetrapod diversity in the Bauru Group is remarkable, including an admixture of typically austral Gondwanan taxa (e.g., abelisaurids, notosuchians) and boreal Gondwanan forms (e.g., carcharodontosaurids). Of note is the absence of Laurasian taxa in the upper portion of the Bauru Group. With the exception of some turtles, an anuran, mesoeucrocodylians and one titanosaur, most taxa from the Bauru Group are based on fragmentary and isolated bones, and as such many specimens can be identified only to a higher taxonomic level. Fishes, turtles, anurans, mesoeucrocodylians, dinosaurs, birds and mammals from the Adamantina and Marília formations resemble the latest Late Cretaceous vertebrate faunas from southern South America, except for the absence of ornithischian dinosaurs.     

Late Cretaceous (Santonian) Atractosteus (Actinopterygii,Lepisosteidae) remains from Hungary (Iharkút,Bakony Mountains)

Lepisosteid fishes are well known from the Upper Cretaceous of Europe, but only by fragmentary remains from some Cenomanian and Campanian–Maastrichtian deposits. Here we report various cranial and postcranial remains of gars, discovered in the Upper Cretaceous (Santonian) Csehbánya Formation of Iharkút (Bakony Mountains, Hungary). These remains represent one of the most diverse assemblages of lepisosteid fish material from Upper Cretaceous continental deposits of Europe. Based on tooth morphology, scale-microstructure and the features of the supracleithrum we refer these remains to the genus Atractosteus. Besides some uncertain remains from the Cenomanian of France and Spain, the Santonian aged fossils from Iharkút represent the oldest undisputable occurrence of the family Lepisosteidae in the European continental Cretaceous. Using tooth crown morphology, the surface microstructure of the ganoid scales and the anatomy of the supracleithrum a review of the Late Cretaceous lepisosteid record suggests the occurrence of both Atractosteus and Lepisosteus in the European archipelago.