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.     

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.     

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.     

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 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.     

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.     

The Early Cretaceous megaflora of the Springhill Formation,Patagonia. Paleofloristic and Paleonvironmental inferences

The Springhill Formation (Lower Cretaceous) is considered as one of the most important oil producer units within the Austral Basin in Argentina. However, knowledge regarding this formation's plant community composition is still very scarce. In this work a thorough survey of the outcrops of the Springhill Formation was conducted regarding both geological and palaeobotanical aspects. Abundant fossil plant material was collected at the Estancia El Salitral and Río Correntoso localities in Santa Cruz Province, Argentina, consisting primarily of compressions of leafy shoots with several branching orders, completely to partially preserved fronds, simple to pinnate leaves and fertile scales. Analysis of these materials made it possible to describe 20 new taxa which add to the already known flora of the Springhill Formation; nine of these taxa were found in Estancia El Salitral, and the remaining 11 were found in Río Correntoso. The floristic diversity of the unit has increased through the discovery of the first megafossils of ginkgophytes (Baiera sp. 1 and Baiera? sp. 2), whereas new taxa have been described within the divisions of Pteridophyta, Pteridospermophyta, Cycadophyta and Pinophyta. Analysis of the vegetative and reproductive epidermal structures of the various groups included within this megaflora has been demonstrated to be a valuable tool for the recognition and differentiation of species. Analysis has also made it possible to suggest paleoenvironmental inferences. Finally, the megaflora from the Springhill Formation was compared to other relevant plant assemblages from the Lower Cretaceous of Western Gondwana. The megaflora was determined to be most similar to that of the well-known Patagonian Anfiteatro de Ticó Formation and most different from that of the Brazilian Crato Formation.     

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.     

Reappraisal of Fresnosaurus drescheri (Plesiosauria; Elasmosauridae) from the Maastrichtian Moreno Formation,California, USA

The elasmosaurid Fresnosaurus drescheri, Welles from the contact between the Tierra Loma/Marca members of the Moreno Formation (Maastrichtian), California, USA is reviewed. Most of the features included in Welles's original diagnosis are considered related only to the juvenile ontogenetic stage of the holotype and only specimen. The new diagnosis is based on diagnostic characters of the ilia, including long rectangular shaped sacral facets located in the dorsal part of the shaft, two gentle knobs in the shafts and unexpanded dorsal end. Additional material from the Moreno Formation (numbered under the same number as the F. drescheri holotype but not mentioned by Welles and therefore considered part of a different specimen) are described for the first time. The latter are referred to the aristonectine, being the first evidence of aristonectines from the North Hemisphere.     

Dinosaur coprolites from the Late Cretaceous (Maastrichtian) Lameta Formation of India: isotopic and other markers suggesting a C3plant diet

A single locality of the Late Cretaceous (Maastrichtian) Lameta Formation at Pisdura in Central India has yielded a large number of coprolites attributed to titanosaurian dinosaurs. Internally the coprolites are dark grey and contain abundant plant tissues and other organic materials. The plant tissues are mostly of gymnospermous origin. In addition, remains of bacterial colonies, fungal spores and algae are seen in the macerated fraction under scanning electron microscope. The dark grey appearance is probably attributable to fine-grained organic matter within voids in tracheids or xylem. The average ¹³C/¹²C ratio of the organic matter in the coprolites is -24‰ (relative to PDB) suggesting that plants of C₃type were the main diet of their producers. A comparison of δ¹⁵N value (about 4‰ w.r.t. air) of the coprolites with that of faecal matter of modern herbivores and carnivores suggests that gut fermentation may not have been an active mechanism in the digestion process of titanosaurs.     

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.     

A rediscovered fossil locality at Shedden,near Burnley,Lancashire

Carbonicola agrestis (Brown 1843), described with C. heyae, C. rugosa and C. smithii, were recorded from “Ironstone Shale” at or near “Sheden”, a locality lost for many years. Non-marine bivalves closely comparable with Brown's figures of C. agrestis and C. heyae were recently discovered in an old tip at Shedden, about 6–5 km south-east of Burnley. With other evidence the material suggests that Brown's species were collected from a small mine in this locality and from a horizon not more than 3.7 m above the Arley Mine.     

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.     

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.     

A new titanosaur (Dinosauria,Sauropoda) from the Upper Cretaceous of Lo Hueco (Cuenca,Spain)

The upper Campanian-lower Maastrichtian site of Lo Hueco (Cuenca, Spain) has provided a set of well-preserved partial skeletons in anatomical connection or with a low dispersion of their skeletal elements. One partial skeleton is herein described and a new titanosaurian sauropod is established, Lohuecotitan pandafilandi. This titanosaur is diagnosed by eight autapomorphic features: dorsally and ventrally widened or bifurcated posterior centrodiapophyseal lamina in anterior and middle dorsal vertebrae; short postspinal lamina with a transversely expanded distal end represented by smooth scars in the dorsal vertebrae; anteriormost caudals with the medial spinoprezygapophyseal and medial spinopostzygapophyseal laminae ventrally connected with the prespinal and postspinal laminae, respectively; anterior caudal neural spines with a dorsal projection of the prespinal and postspinal laminae; anterior caudal neural spines bears a “greek-cross”-like cross-section; middle caudal centra having two round and rough structures in the dorsal edge of the posterior articulation, which extends to the dorsal surface of the centrum; the articular ends of the rami of the haemal arches are divided in two articular surfaces; and tuberosity between the anterior and the lateral trochanter of the fibula. The herein performed phylogenetic analysis considered L. pandafilandi as a member of Lithostrotia more derived than Malawisaurus. The known palaeodiversity of the Late Cretaceous Ibero-Armorican titanosaurs is increasing, and further analyses focused on this group will be necessary to better understand the evolutionary history of European titanosaurs and to clarify their relationships within Titanosauria.     

Palaeomagnetic,geochronological and geological constraints on the tectonic evolution of the Mejillones Peninsula,northern Chile

Palaeomagnetic and geochronological data from an Early Palaeozoic high grade metamorphic complex (Jorgina Formation) and Jurassic layered basic intrusion (Moreno Complex) are reported from the Mejillones Peninsula of northern Chile (23–23°30'S). ⁴⁰Ar–³⁹Ar dates from the Lower Palaeozoic Jorgina Formation and the Moreno Complex are between 170 and 158 Ma, coincident with a phase of emplacement of the north Chilean coastal batholith. This suggests that intrusion and magnetization of the Moreno Complex and the metamorphism and remagnetization of the Jorgina Formation were related to batholith emplacement. Extracted stable components of magnetization from all units (17 sites) define site-mean directions with a scattered distribution. The scatter in site-mean directions is interpreted as being due to minor, localized, non-uniform, block-fault related (normal or strike-slip, or both) rotation after 158 Ma. The palaeomagnetic and geochronological data indicate that no significant large-scale latitudinal translation of crustal blocks has taken place in this part of northern Chile since the Late Jurassic. In addition, they indicate that the uniform clockwise rotation after the mid-Cretaceous which affected the adjacent Cordillera de la Costa either did not extend into the Mejillones Peninsula or took the form of localized block-fault rotations. The restriction of palaeomagnetically defined styles of rotation to discrete areas within the north Chilean forearc indicates that forearc wide block-fault rotation models are not applicable to the Pacific margin of northern Chile.     

Carbonaceous filaments from North Pole,Western Australia: Are they fossil bacteria in Archaean stromatolites?

A sample of chert from North Pole in the Archaean Pilbara block of Western Australia contains carbonaceous filaments that resemble microfossils. These occur in alternating light and dark laminae that look stromatolitic. However, the filaments are too simple in form for their origin to be determined, so they should be regarded as dubiofossils, perhaps biogenic, perhaps inorganic. Their host laminae were inorganically precipitated in a concordant fissure and thus cannot be stromatolitic. This fissure is younger than the surrounding silicified sediments of the ca. 3500 Ma old Warrawoona Group and possibly formed towards the end of the uplift and associated fracturing of the North Pole Dome, perhaps ca. 2750 Ma ago. The filaments are therefore contaminants in secondary chert.The filament-bearing rock was collected less than a metre from one of the localities (B) from which Awramik et al. reported early Archaean microfossils and possible microfossils. Their filaments from this locality were almost identical to those described here and were found in similar laminae. This suggests that their locality B filaments may also be contaminants in secondary chert. Other filaments found by Awramik et al. at North Pole come from an imprecisely located sample site (locality A) where the rock relationships are unknown. Since the host laminae of these filaments are not demonstrably primary and as cryptic concordant fissures filled with secondary minerals are common in locality A rocks, the filaments from this sample site could be contaminants too. Those that were assigned to Archaeotrichion should be treated as dubiofossils. Thus, the filaments described by Awramik et al. may not be fossil bacteria in ca. 3500 Ma old stromatolites, as they proposed, and are not necessarily the oldest known fossil organisms, as has been claimed.     

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.     

Facies architecture of a Lower Cretaceous coral-rudist patch reef, Arizona

The Lower Cretaceous Mural Limestone marks the maximum marine incursion into southeast Arizona during Aptian-Albian time and records the middle Cretaceous transition from coral-dominated to rudist-bivalve dominated reefs. Upper Mural Limestone facies are most often dominated by corals. However, rudists form significant frameworks at some localities, one of which is described in this paper. The paleoenvironmental distribution of three potential reef-builders (corals, rudists, and ‘oysters’) were studied at this patch reef locality. Corals built the framework of the inner reef core. The rudist Petalodontia initially gained a foothold in sheltered areas among corals and subsequently built a framework in the outer reef core. Caprinid rudists formed mounds in the outer reef to back reef areas. The rudists Toucasia and Monopleura and the oyster-like bivalve Chondrodonta formed beds or were scattered in the reef-flank and shelf lagoon sediments and did not contribute to the reef framework.Upper Mural Limestone reefs are important examples of the coexistence of corals and rudists during this middle Cretaceous faunal transition period. This study supports the idea that rudist-bivalves initially colonized protected back-reef areas early in the Cretaceous and only later in the Cretaceous did rudists dominate reef frameworks.     

Occurrence of angiosperm remains in an Early Cretaceous intertrappean bed, Rajmahal Basin, India

An angiosperm fruit of uncertain affinity and a questionable flower are reported from the Rajmahal Formation (Early Cretaceous) at the locality of Sonajori, Rajmahal Basin. The remains of ferns, Pentoxylales and conifers have been described previously from this locality. The significant addition of angiosperm megafossil remains confirms that flowering plants were evolving during the Early Cretaceous Epoch in India. The Sonajori assemblage seems to be the youngest fossil assemblage recovered so far from the Rajmahal Basin. It is tentatively dated as Barremian–Aptian.