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This paper reports the use of FEA (Finite Element Analysis) to model dinosaur tracks. Satisfactory reproductions of sauropod ichnites were simulated using 3D numerical models of the elasto-plastic behaviour of soils. Though the modelling was done of ichnites in situ at the Miraflores I tracksite (Soria, Spain), the methodology could be applied to other tracksites to improve their ichnological interpretation and better understand how the type and state of the trodden sediment at the moment the track is created is a fundamental determinant of the morphology of the ichnite. The results obtained explain why the initial and commonly adopted hypothesis—that soft sediments become progressively more rigid and resistant at depth—is not appropriate at this tracksite. We explain why it is essential to consider a more rigid superficial layer (caused by desiccation) overlying a softer layer that is extruded to form a displacement rim. Adult sauropods left trackways behind them. These tracks could be filled up with water due to phreatic level was close to the ground surface. The simulation provides us with a means to explain the differences between similar tracks (of different depths; with or without displacement rims) in the various stratigraphic layers of the tracksite and to explain why temporary and variable conditions of humidity lead to these differences in the tracks. The simulations also demonstrate that track depth alone is insufficient to differentiate true tracks from undertracks and that other discrimination criteria need to be taken into account. The scarcity of baby sauropod tracks is explained because they are shallow and easily eroded.
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The Galinha tracksite reveals a sequence of Bajocian–Bathonian limestones belonging to the Serra de Aire Formation (West−Central Portugal) and is one of the few sites in the world where Middle Jurassic sauropod dinosaur tracks can be found. This tracksite is characterised by the presence of long, wide gauge sauropod trackways, the Middle Jurassic age of which suggests these dinosaurs were more widely distributed over time than previously thought. Two trackways contain unique pes and manus prints with morphologies that allow a new sauropod ichnotaxon to be described: Polyonyx gomesi igen. et isp. nov. On the basis of different manus/pes prints and trackway features, the proposal is made to subdivide Sauropodomorpha ichno−morphotypes into five groups: Tetrasauropus−like, Otozoum−like, Breviparopus/Parabrontopodus−like; Brontopodus−like, and Polyonyx−like. Polyonyx gomesi igen. et isp. nov. is thought to represent a nonneosauropod eusauropod, with a well developed manus digit I. The posterior orientation of this digit print suggests they were made by a eusauropod dinosaur with a posteriorly rotated pollex. The manus print morphologies observed in two trackways suggest a stage of manus structure intermediate between the primitive non−tubular sauropod manus and the tubular metacarpal arrangement characteristic of more derived sauropods. The low heteropody (manus:pes area ratio 1:2) of the trackway renders it possible they could have been made by eusauropods such as Turiasaurus riodevensis, which has a similar manus:pes area ratio. The Polyonyx igen. nov. trackway was made by non−neosauropod eusauropod, and suggests that wide gauge sauropod trackways were not exclusively made by Titanosauriformes.
Neuquensaurus, from the Late Cretaceous of Argentina and one of the first dinosaurs described from Patagonia, is one of the most derived sauropod dinosaurs, and its proportions and size place it among the smallest sauropods ever known. In this context, Neuquensaurus is central to understanding late stages of sauropod evolution. This contribution offers a full description of the appendicular skeleton of Neuquensaurus. The anatomical analysis reveals that the appendicular skeleton of Neuquensaurus exhibits unique characteristics only shared with closely related saltasaurine titanosaurs; for example, the laterally directed preacetabular lobe of the ilium, the prominent fibular lateral tuberosity, and the presence of an intermuscular line on the femoral shaft, which is proposed here as a synapomorphy of Saltasaurinae. Neuquensaurus also displays many reversals to primitive character states, such as the presence of a prominent olecranon process of the ulna, a trochanteric shelf, a lesser trochanter and an ischial tuberosity. Additional characters that allow its evaluation in a phylogenetic context are here provided. Among them are the extremely deflected femoral shaft, the elliptical femoral cross−section, and the anterolaterally oriented cnemial crest.
A new medium−sized rebbachisaurid sauropod from the Castrillo la Reina Formation (Upper Barremian–Lower Aptian) in Burgos Province, Demandasaurus darwini gen. et sp. nov., is described. It is known from an incomplete but associated skeleton that includes cranial and post−cranial remains. Demandasaurus darwini gen. et sp. nov. presents 9 autapomorphies in the teeth and vertebrae. Demandasaurus is the first diplodocoid sauropod described from the Cretaceous of the Iberian Peninsula. Its inclusion in the Rebbachisauridae is well supported by our phylogenetic hypothesis, which situates it as a sister group of Nigersaurus from the Aptian of Niger, with which it shares various synapomorphies. The discovery of Demandasaurus provides further evidence of the sporadic use of the Apulian Route by dinosaurs during the Early Cretaceous for moving between the south of Europe (Laurasia) and the north of Africa (Gondwana).
Lirainosaurus is the only titanosaurian sauropod described to date from the Late Cretaceous of the Iberian Peninsula. The type of Lirainosaurus astibiae consists of both cranial and postcranial remains that were found as disarticulated elements in the Laño quarry (Treviño, northern Spain). This taxon was diagnosed originally on the basis of vertebral and appendicular autapomorphic traits. The study of a paratypic skull fragment and a second referred specimen provides information about its braincase morphology. Lirainosaurus is regarded as a derived titanosaur on the basis of the complete fusion between the prootic and the exoccipital−opisthotic complex, the position of the cranial foramina, and the shape and orientation of the occipital condyle. The braincase of L. astibiae appears to be diagnostic in the presence of a foramen distally on each basal tubera. The absence of median subcondylar foramina in the basioccipital may be an autopomorphic trait or be due to ontogenetic growth. A comparison with other partial skulls known in Europe suggests a high diversity during the Campanian/Maastrichtian, with at least three different titanosaurian species living in the Ibero−Armorican Island.
The stegosaurian forelimb is usually portrayed with the metacarpals slanted and distally spread. However, manual manipulation of stegosaurian metacarpals reveals that in that configuration they do not articulate with each other nor with the rest of the forelimb. Rather, they do articulate with each other and with the rest of the forelimb when posed vertically and arranged in a compact, semi−tubular configuration, as in sauropods. This configuration agrees with data from articulated specimens and trackways. As with sauropods, this metacarpal configuration makes retention of phalanges awkward for locomotion and may be functionally related to the vestigiality of the manual phalanges of the outer digits.
Knowledge of titanosaurian cranial anatomy has improved substantially in the last decade because several skulls have come to light or were restudied. The discovery of Bonitasaura salgadoi, a partial titanosaurian skeleton including cranial bones, permitted the definitive recognition of square jaws in a titanosaurian sauropod as well as a peculiar skull morphology that increases the morphological diversity of the group. Here we present a full description and illustration of the skull material of B. salgadoi. Among cranial bones, the lacrimal, quadrate, and dentary exhibit apomorphic differences from those of other titanosaurians. Conversely, the frontal and parietal are more conservative. A phylogenetic analysis recovers B. salgadoi as a member of the Titanosauria, related to mid−sized to large titanosauroids from the Turonian–Campanian of South America, in contrast to a previous hypothesis that suggested a nemegtosaurid affinity. The skull reconstruction presented here shows that the skull of B. salgadoi is anteroposteriorly short and dorsoventrally high, contrasting with the elongate skull of Rapetosaurus krausei.
The dinosaur record of the Salitral Moreno locality (Río Negro Province, Argentina) is characterized by a high diversity of herbivore taxa, among them hadrosaurs, ankylosaurs, and titanosaur sauropods, but carnivores are rare, consisting of only a few fragmentary bones of small forms. Titanosaurs are represented by Rocasaurus muniozi and Aeolosaurus sp., and at least four other taxa, represented by fragmentary material. The elements preserved include a cervical, dorsal and caudal vertebrae, chevron, humerii, ulnae, radii, metacarpal, femora, tibiae, metatarsal, ischia, pubis, and ilium. The Allen Formation is thought to be correlated with the Marília Formation in Brazil, and their faunas have certain elements in common such as aeolosaurines, but saltasaurines and hadrosaurs, are known exclusively from the Allen Formation. These absences, and particularly that of the saltasaurines, may be because those sauropods originated late in the Cretaceous, probably in southern South America (Northern Patagonia?), and they did not have time to disperse to northern South America.
Sauroposeidon proteles is a large brachiosaurid sauropod recently described from the Antlers Formation (Aptian-Albian) of southeastern Oklahoma. Sauroposeidon represents the culmination of brachiosaurid trends toward lengthening and lightening the neck, and its cervical vertebrae are characterized by extensive pneumatic structures. The elaboration of vertebral air sacs during sauropod evolution produced a variety of internal structure types. We propose a new classification system for this array of vertebral characters, using computed tomography (CT) of pneumatic internal structures. Comparisons with birds suggest that the vertebrae of sauropods were pneumatized by a complex system of air sacs in the thorax and abdomen. The presence of a thoraco-abdominal air sac system in sauropods would dramatically affect current estimates of mass, food intake, and respiratory requirements. Sauroposeidon was one of the last sauropods in the Early Cretaceous of North America; sauropods disappeared from the continent by the early Cenomanian. The demise of sauropods in the Early Cretaceous of North America predates significant radiations of angiosperms, so the decline and extinction of this dinosaur group cannot be linked to changes in flora.
Exceptionally well−preserved embryonic skulls of Upper Cretaceous (Campanian Anacleto Formation) sauropods from Auca Mahuevo (Neuquén Province, Argentina) provide important insights into the ontogeny and evolution of titanosaurian neosauropods. The most important cranial modifications occurring during titanosaurian ontogeny appear to be centered on the infraorbital and narial regions, which exhibit a substantial degree of “mosaic” evolution. On one hand, the Auca Mahuevo embryos show a large jugal that forms part of the lower margin of the skull and unretracted external nares, as indicated by the position and orientation of the lacrimals as well as the anterior extension of the frontals. Both of these features are ancestral for neosauropods, being present in prosauropods. On the other hand, the embryonic skull exhibits a large ventral notch, tentatively interpreted as homologous to the neosauropod preantorbital fenestra, that opens ventral to the jugal and between the maxilla and the quadratojugal, and a temporal region that closely resembles the adult neosauropod condition. This mosaic of character states indicates that different regions of the skull of titanosaurian neosauropods acquired their characteristic morphology at substantially different rates during their ontogenetic development.
The axial soft−tissue system in the neck of Dicraeosauridae and Diplodocidae, including pneumatic diverticula, ligaments, and muscles, is reconstructed on the basis of phylogenetic and functional morphological comparisons with extant crocodylians and birds and compared with other soft−tissue reconstructions for sauropods. Bifurcation of the neural spines separated the paired supraspinal ligament into two sheets. A paired interspinal septum was attached to the cranial and caudal margins of the neural spines. The dorsal and the lateral portions of the cervical musculature must have been strongly segmented, whereas the laterocostal portion was divided with one myoseptum per vertebral segment. The hypaxial cervical muscle was most probably small and only poorly segmented. In Diplodocidae and Dicraeosauridae, the distribution of external pneumatic structures is similar, whereas only Diplodocidae possess intraosseous pneumatic structures. Supravertebral pneumatic diverticula are reconstructed for both groups, which, together with dorsal ligaments filled the gap between the metapophyses of bifurcate neural spines. Comparisons between the vertebrae of juvenile and adult diplodocids strongly indicate that pneumatisation proceeded from the supramedullary diverticula into the neural arch and the neural spine. The regular branching pattern of the pneumatic cavities as well as the vertical I−beam construction of the vertebral corpora is interpreted as a consequence of the biomechanical constraints of the vertebral corpora in diplodocids. These reconstructions form the ground for functional morphological considerations in Diplodocidae and Dicraeosauridae while addressing the possible mechanical consequences of pneumatic structures for the integrity of the support system of the neck.
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