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 Titanosaurian Sauropod Dinosaur with Opisthocoelous Caudal Vertebrae from the Early Late Cretaceous of Liaoning Province, China

We describe a new titanosaurian sauropod dinosaur, Borealosaurus wimani gen. et sp. nov., based on a distinctive mid-distal caudal vertebra from the early Late Cretaceous Sunjiawan Formation exposed in the Shuangmiao village of Beipiao in Liaoning, China. We provisionally refer an isolated tooth crown, a middle caudal vertebra, and a right humerus from the same locality and horizon to this taxon. Borealosaurus is distinguished from other sauropods in its possession of opisthocoelous mid-distal caudal vertebrae. The occurrence of opisthocoelous caudals in Borealosaurus and the Mongolian sauropod Opisthocoelicaudia raises the possibility that these taxa pertain to an as-yet unrecognized titanosaurian subclade endemic to the Cretaceous Asia.     

Description of a juvenile titanosaurian dinosaur from the Upper Cretaceous of Brazil

We describe a juvenile specimen of a titanosaurian sauropod dinosaur consisting of two dorsal and three caudal vertebral centra, an ilium fragment, and an ischium unearthed in 1991 from Site Km 153.5 at BR-050 highway in the Serra da Galga region, municipality of Uberaba, state of Minas Gerais, Brazil. The outcrop at the site is assigned to the Serra da Galga Member of the Marília Formation (Bauru Basin: Bauru Group; Upper Cretaceous: Maastrichtian). Although the material is very incomplete, features such as strongly procoelous caudal centra suggest an affinity with the titanosaurian clade Lithostrotia. The extensive vertebral pneumaticity with deep pleurocoels and well-developed camerae supports the hypothesis that, in titanosaurs, the air sac system was already present and fully developed even at early ontogenetic stages.     

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.     

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 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.     

Caudal pneumaticity in derived titanosaurs (Dinosauria: Sauropoda)

Among extant vertebrates, only birds have a respiratory system associated with pneumatic diverticula. However, several extinct clades also had pneumatic diverticula, including the sauropod dinosaurs. Among sauropods, Saltasaurini are characterized by extreme postcranial pneumaticity, which extends as far down the spinal column as the posterior caudal vertebrae. In this paper the pneumatic foramina in both the neural arches and the centra are described in detail, and the relative proportion of air spaces in the caudal vertebrae is established, revealing that the skeleton of Rocasaurus muniozi was more pneumatized than that of Neuquensaurus australis, with Saltasaurus loricatus intermediate. The level of pneumatization varies between the three saltasaurine taxa: in Neuquensaurus, only the neural arch is pneumatized, whereas in the other two saltasaurines both neural arches and centra are pneumatized. This allows us to hypothesize that the timing of pneumatization varied between the three species, with Rocasaurus muniozi pneumatized earliest in ontogeny. This ontogenetic pattern is correlated with evolutionary derivation: the most derived taxa show pneumatization in both the neural arch and the centrum.     

New materials of lithostrotian titanosaurs (Dinosauria: Sauropoda) from the Upper Cretaceous of central Patagonia

Continental Cretaceous deposits exposed in central Patagonia, Argentina, preserve a rich and important record of titanosaurian evolution that spans the entire Upper Cretaceous. Recently, a new lithostragraphic unit, the Lago Colhué Huapi Formation (Coniacian – Maastrichtian), was located in the Golfo San Jorge Basin. Here, we describe new titanosaurian sauropod material from that formation (UNPSJB-Pv 1051). The material consists of a partially articulated left hind limb and disarticulated but associated skeletal elements. They are confidently referred to Titanosauria, and within that clade, to Lithostrotia, probably occupying a derived position. We interpret the bone concentration as an assemblage of hydraulic origin deposited in a fluvial channel. This new material begins the incipient fossil record of the Lago Colhué Huapi Formation, thereby also increasing the titanosaurian diversity of the latest Cretaceous. Additionally, the new titanosaurian enhances evidence in support of strong faunal similarities among the Allen and Marilia formations, chronological equivalents in the Cretaceous of South America. Furthermore, the Lago Colhué Huapi specimen adds to the Late Cretaceous record of Titanosauria and augments our knowledge of central Patagonian terrestrial vertebrate assemblages during this interval.     

The axial skeleton of the titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from the latest Cretaceous of Spain

Lirainosaurus astibiae was originally described by Sanz and collaborators in 1999 on the basis of a skull fragment, isolated teeth, several vertebrae (e.g. the holotypic anterior caudal vertebra) and appendicular bones from the Late Cretaceous of Laño (northern Spain). A review of all the vertebral remains, including new material (cervical, dorsal and caudal vertebrae, dorsal ribs, haemal arch), provides additional information about the axial skeleton of Lirainosaurus. A study of the laminae and fossae shows interesting variations in these structures in the axial series, especially concerning the prezygapophyses and diapophyses: e.g. the X-shaped morphology of the centroprezygapophyseal lamina only in the posterior dorsal vertebrae, and the division of the postzygodiapophyseal fossa into two fossae in the posterior dorsal vertebrae and the proximal caudal vertebrae. Two vertebral characters are here considered to be autapomorphic for L. astibiae: the presence of a lamina in the interzygapophyseal fossa in the most proximal caudal vertebrae (a postzygodiapophyseal lamina that separates the ventral postzygapophyseal centrodiapophyseal fossa and the dorsal postzygapophyseal spinodiapophyseal fossa), and the spinopostzygapophyseal structure not posteriorly projected in the posterior caudal vertebrae. The combination of characters present in the axial remains of Lirainosaurus astibiae supports the idea that it is a derived lithostrotian close to Saltasaurinae.     

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.