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.     

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.     

Osteoderms of Montealtosuchus arrudacamposi (Crocodyliformes,Peirosauridae) from the Turonian-Santonian (Upper Cretaceous) of Bauru Basin,Brazil

The skin of Crocodyliformes is characterized by osteoderms of various sizes and shapes. It plays roles in defence, thermoregulation, sexual attraction, calcium reserves, and locomotion. This study presents the morphometric characteristics of osteoderms preserved in the nuchal, dorsal, ventral and appendicular shield of Montealtosuchus arrudacamposi, a Peirosauridae crocodyliform from the Turonian-Santonian (Upper Cretaceous) of Adamantina Formation, Bauru Basin, Brazil. The results of the analysis show that its dermal shield had protective and thermoregulatory functions and a well-defined mechanical function allowing terrestrial locomotion and enabling good agility and capacity for movement.     

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 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.     

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.     

New record of titanosaurian (Dinosauria: Sauropodomorpha) osteoderms from the Upper Cretaceous of North Patagonia

The idea that titanosaurs had osteoderms was proposed in the late nineteenth century by Depéret. However, this idea was given little credence by other researchers until 1980 when unequivocal evidence of armoured titanosaurs was reported. Since then, many discoveries of titanosaurian osteoderms have been made worldwide. In this work, seven osteoderms are described from the Allen Formation (upper Campanian–lower Maastrichtian), Salitral Moreno locality, Río Negro Province, Argentina. Among the described osteoderms it was possible to recognize three morphologies (keeled, ellipsoidal and cylindrical), with the first of these being most prevalent. Although the osteoderms from Salitral Moreno resemble osteoderms found in other parts of the world, no osteoderms of a similar morphotype are known from this locality, or from Lago Pellegrini or Cinco Saltos.     

Cranial anatomy of Pholidosaurus purbeckensis from the Lower Cretaceous of France and its bearing on pholidosaurid affinities

New and nearly complete cranial remains of Pholidosaurus purbeckensis are described on the basis of specimens recovered from the Berriasian locality of Cherves-de-Cognac, France. Two skulls, associated mandibles and a set of dorsal osteoderms are available and allow a refinement of the anatomy of the genus, known otherwise from coeval deposits in Germany and England. Because of its longirostrine morphology, convergent with other crocodylomorph lineages, the phylogenetic relationships of Pholidosaurus are likely to be affected by Long Branch Attraction problems. Various tests of removing/excluding longirostrine lineages confirm that Dyrosauridae have a labile position and that their affinities with Pholidosauridae are weakly supported. Results from comparative anatomy and phylogenetic analyses recover Pholidosaurus as the basalmost member of Pholidosauridae, a group closely related to Goniopholididae. Pholidosaurus is recovered together with abundant remains of Goniopholis and with more limited remains of Theriosuchus. The paleoecology of Pholidosaurus and more generally, of Pholidosauridae, is discussed.     

First sauropod bones from Italy offer new insights on the radiation of Titanosauria between Africa and Europe

Here we describe the first sauropod skeletal remains from the Italian peninsula that also represent the earliest record of titanosaurs in Southern Europe. Scattered bones, including an almost complete anterior caudal vertebra, were found in Cretaceous (Aptian–Albian) marine deposits, some 50 km East of Rome. The vertebra shows a bizarre and perhaps unique orientation of the zygapophyseal articular facets that renders their interpretation problematic. Phylogenetic retrofitting tests support the placement of the Italian titanosaur among basal lithostrotians. Palaeobiogeographic analysis based on the resulting phyletic relationships suggests an Afro-Eurasian route for the ancestors of the Italian titanosaur, a scenario compatible with the palaeogeographic evolution of the Italian microplates during the Cretaceous. Together with previously recorded titanosaurian-like ichnites from a Cenomanian locality in Latium, this new find suggests a quite long emersion for the Apenninic carbonate platform. We suggest that the Italian titanosaur was member of a population that crossed the western Tethys Sea through a “filtering bridge” composed of a chain of ephemeral islands and peninsulae, known as Periadriatic (Adria) carbonate platforms, that connected sporadically Africa and Europe since the Early Cretaceous.     

Late Pliocene Glyptodontinae (Xenarthra,Cingulata, Glyptodontidae) of South and North America: Morphology and paleobiogeographical implications in the GABI

Knowledge of the main aspects of the Great American Biotic Interchange (GABI) concerning the glyptodontine Glyptodontidae (Xenarthra) is very scarce. A bidirectional dispersal process was recently proposed for this clade, with the presence of the North American genus Glyptotherium Osborn recognized in latest Pleistocene sediments of northern South America (Venezuela and Brazil). However, the earliest stages of this paleobiogeographical process remain poorly understood, mainly because of the limited fossil record on this clade in late Pliocene sediments. The goals of this contribution are: a) to present and describe the first record of a glyptodontine glyptodontid from the late Pliocene of northern South America, tentatively assigned to a new species of Boreostemma Carlini et al. (Boreostemma? sp. nov); and b) to analyze its paleobiogeographical implications with respect to the GABI. This new material was recovered from the San Gregorio Formation (late Pliocene, prior the GABI) in northern Venezuela, where it is represented by several osteoderms of the dorsal carapace. A comparison among the three known late Pliocene glyptodontine glyptodontids of a) southern South America (Paraglyptodon), b) northern South America (Boreostemma), and c) southern North America ("Glyptotherium"), reveals a series of shared characters between (b) and (c), not present in (a). The most important of these shared characters in (b) and (c) are: all the osteoderms present a great development of the central figure, which is always larger than the peripherals; the sulcus that delimits the central and peripheral figures is narrower and shallower; and all the osteoderms present are relatively thin. This evidence suggests that the lineage of Glyptodontinae which participated in the GABI and subsequently diversified in North America originated in northern South America. Moreover, the evident morphological differences between these glyptodontines with respect to the southern South American forms show a significant separation of both lineages since at least latest Miocene-early Pliocene.     

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.     

Lower Cretaceous Mesoeucrocodylians from Scandinavia (Denmark and Sweden)

The crocodyliform faunas of the lowermost Cretaceous Rabekke and Jydegård Formations on the Baltic island of Bornholm, Denmark, and the Annero Formation of Skåne, southernmost Sweden, are represented by isolated teeth, osteoderms, and vertebrae. The rich Berriasian assemblage of the Rabekke Formation includes at least three distinctive taxa: Bernissartia sp., Theriosuchus sp., and Goniopholis sp., an association that is also known from several other contemporaneous European vertebrate localities. In contrast to this fauna, the Jydegård and Annero Formations have yielded only rare mesoeucrocodylian remains, which are assigned to Theriosuchus sp. and an undetermined mesoeucrocodylian taxon, possibly Pholidosaurus. Geographically, the Scandinavian localities represent the easternmost and northernmost distribution of typical continental Jurassic-Cretaceous crocodyliform communities in Europe.     

A saltasaurine titanosaur (Sauropoda: Titanosauriformes) from the Angostura Colorada Formation (upper Campanian,Cretaceous) of northwestern Patagonia,Argentina

Saltasaurine titanosaurs are characterized by their relatively small size compared to other sauropods, extreme postcranial pneumaticity, and dermal armour covering the body. This group has been reported in the Upper Cretaceous of the Lecho, Allen, and Anacleto formations of Argentina. We describe here a new saltasaurine specimen (MACN-Pv RN 233) from the Campanian of the Angostura Colorada Formation (Río Negro Province) that is represented by eight caudal vertebrae and six osteoderms. This specimen is described in detail and its phylogenetic relationships with the other three known saltasaurines, as well as its implications to the knowledge of caudal vertebra and osteoderm anatomy, are discussed. Our results place MACN-Pv RN 233 more closely related to Saltasaurus loricatus and Rocasaurus muniozi than to Neuquensaurus australis. MACN-Pv RN 233 possesses a combination of features that differ from other saltasaurines, but because of the fragmentary nature of the specimen we decided for the sake of taxonomic stability to not erect a new taxon. This specimen shows the first unambiguous evidence of chevron pneumatisation for a sauropodomorph, implying a broader osteological invasion of the diverticula from the abdominal air sac than previously thought for this group of dinosaurs. MACN-Pv RN 233 preserves two osteoderm morphotypes, one similar to those reported for Neuquensaurus australis and Saltasaurus loricatus. This new specimen expands the distribution of the group to a new geological unit and increases the dinosaur diversity known for the Angostura Colorada Formation.     

Internal vascularity of the dermal plates of Stegosaurus (Ornithischia, Thyreophora)

X-ray computed tomography and petrographic thin sectioning were used to study internal features of the plates of the thyreophoran dinosaur Stegosaurus and the osteoderms of Alligator. Infrared thermographic imaging of basking caimans was used to examine possible differential blood flow to osteoderms and other parts of the skin. Multiple large openings in the Stegosaurus plate base lead to a linear, mesiodistally oriented vestibule, which in turn apically sends off multiply branching “pipes”. The pipes are best developed in the basal half of the plate, and communicate with cancellous regions (some of which presumably were vascular spaces) throughout the plate interior. Some internal vascular features also connect with vascular pits and grooves on the plate surface. Alligator osteoderms show a similar internal vascularity. In crocodylians, the osteoderms serve as armor and help to stiffen the back for terrestrial locomotion, but their vascularity enables them to be used as sources of calcium for egg shelling, as sites of lactate sequestration, and possibly for heat exchange with the external environment, as suggested by our infrared thermographic imaging of basking caimans. Thyreophoran osteoderms presumably had multiple functions as well. In Stegosaurus the potential thermoregulatory role of the plates may have been greater than in other thyreophorans, by virtue of their extensive external and internal vascularity, their large size, thin cross-sections above the plate base, dorsal position, and alternating arrangement.     

Geological and Palaeontological context of three new Barremian (Lower Cretaceous) vertebrate sites in the Iberian Peninsula (Cuenca Province,Central Spain)

Three new Lower Cretaceous vertebrate sites (Vadillos-1, Vadillos-2, El Tobar) have been recently discovered and studied in the Cuenca Province (Central Spain). They are located in deposits of “Wealden” facies belonging to the El Collado Sandstone and Clay Formation. In these outcrops, micro and macroremains corresponding to plants, invertebrates and vertebrates have been collected and subsequently assigned to macrophytes, charophytes (e.g., Atopochara trivolvis triquetra, Globator maillardii trochiliscoides, Clavator harrisii harrisii), ostracods (e.g., Cypridea gr. modesta, Cypridea cf. C. isasae, Cypridea sp. aff. C. moneta, Cypridea sp. 1, Cypridea sp. 2), molluscs (Unionoida, Viviparus sp.), fishes, amphibians, turtles (cf. Eucryptodira), crocodyliforms (Neosuchia) and dinosaurs (ankylosaurs, ornithopods, theropods). Among the vertebrate remains, scales, teeth, plates, osteoderms, phalanges, ribs, vertebrae and other incomplete bones, as well as eggshell fragments have been identified. This rich and diverse assemblage was deposited in an upper Barremian alluvial-palustrine muddy floodplain crossed by braided sandy channels.     

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.     

Paleocene resurrection of a crocodylomorph taxon: Biotic crises,climatic and sea level fluctuations

Postcranial material of a crocodylomorph from the Danian of Oulad Abdoun Basin (Morocco) is described. Several characters, in particular the shape of its dorsal osteoderms, allows its attribution to Pholidosauridae. Up till now, the latest known pholidosaurid was Terminonaris, from the Early Turonian of North America, so, the Moroccan pholidosaurid extends the stratigraphic range of the group to more than 20 million years younger, and shows that the pholidosaurids survived the K-Pg crisis. The reevaluation and the phylogenetic analysis of Dakotasuchus kingi, Woodbinesuchus byersmauricei, and Sabinosuchus coahuilensis, previously considered as goniopholidids and dyrosaurid, respectively, revealed rather that they are pholidosaurids. This analysis also suggests that at least two independent pholidosaurid lineages reached the Maastrichtian, among which one crossed the K-Pg boundary. This study proposes the first analysis of tethysuchian diversity from the Late Jurassic to the Early Paleogene. Two diversity peaks are observed during the Oxfordian and Cenomanian, two stages of high paleotemperatures evaluated with the δ¹⁸O. The tethysuchian diversity strongly decreases after the Cenomanian, a decline that may be correlated with the “Ocean Anoxic Event” (OAE 2), which caused the strong marine faunal turnover during the Cenomanian-Turonian time interval. The large Turonian-Coniacian oceanic regression could also be a factor in the tethysuchian decline. After that, tethysuchian diversity remains low until the Maastrichtian-lower Paleocene marine tethysuchian dyrosaurid diversification, correlated with the Late Cretaceous-Paleocene transgression. At least one pholidosaurid lineage crossed the K-Pg boundary, confirming the weak impact of this crisis on the crocodylomorphs. Pholidosaurids seem to go extinct at the beginning of the Paleocene, possibly due to the strong radiation of dyrosaurids in marine environments and crocodylians in fresh-water during this time period.