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The recurrent laryngeal nerve is an often cited example of “unintelligent design” in biology, especially in the giraffe. The nerve appears early in embryonic development, before the pharyngeal and aortic arches are separated by the development of the neck. The recurrent course of the nerve from the brain, around the great vessels, to the larynx, is shared by all extant tetrapods. Therefore we may infer that the recurrent laryngeal nerve was present in extinct tetrapods, had the same developmental origin, and followed the same course. The longest−necked animals of all time were the extinct sauropod dinosaurs, some of which had necks 14 meters long. In these animals, the neurons that comprised the recurrent laryngeal nerve were at least 28 meters long. Still longer neurons may have spanned the distance from the end of the tail to the brainstem, as in all extant vertebrates. In the longest sauropods these neurons may have been 40–50 meters long, probably the longest cells in the history of life.
Postcranial skeletal pneumaticity (PSP) is present in a range of basal sauropodomorphs spanning the basal sauropodomorph–sauropod transition. We describe the PSP of five taxa, Plateosaurus engelhardti, Eucnemesaurus fortis, Aardonyx celestae, Antetonitrus ingenipes, and an unnamed basal sauropod from Spion Kop, South Africa (hereafter referred to as the Spion Kop sauropod). The PSP of Plateosaurus is apparently sporadic in its occurrence and has only been observed in very few specimens, in which it is of very limited extent, affecting only the posterior cervical vertebrae and possibly the mid dorsals in one specimen. The PSP of Eucnemesaurus, Aardonyx, Antetonitrus, and the Spion Kop sauropod consists of subfossae (fossa−within−fossa structures) that excavate the vertices of the posterior infradiapophyseal fossae of the posterior dorsal vertebrae. These subfossae range from simple shallow depressions (Eucnemesaurus) to deep, steepsided, internally subdivided and asymmetrically developed chambers (Antetonitrus). The middle and anterior dorsal vertebrae of these taxa lack PSP, demonstrating that abdominal air sacs were the source of the invasive diverticula. The presence of pneumatic features within the infradiapophyseal fossae suggest that the homologous fossae of more basal saurischians and dinosauriforms were receptacles that housed pneumatic diverticula. We suggest that it is probable that rigid non−compliant lungs ventilated by compliant posterior air sacs evolved prior to the origination of Dinosauria.
The neck posture of sauropod dinosaurs has long been controversial. Recent reconstructions position the cervical vertebrae and skull in an “osteological neutral pose” (ONP), the best fit arrived at by articulating the vertebrae with the zygapophyses in maximum contact. This approach in isolation suggests that most or all sauropods held their necks horizontally. However, a substantial literature on extant amniotes (mammals, turtles, squamates, crocodilians and birds) shows that living animals do not habitually maintain their necks in ONP. Instead, the neck is maximally extended and the head is maximally flexed, so that the mid−cervical region is near vertical. Unless sauropods behaved differently from all extant amniote groups, they must have habitually held their necks extended and their heads flexed. The life orientation of the heads of sauropods has been inferred from the inclination of the semi−circular canals. However, extant animals show wide variation in inclination of the “horizontal” semi−circular canal: the orientation of this structure is not tightly constrained and can give only a general idea of the life posture of extinct animals’ heads.
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.
We describe a reasonably complete sauropod foot from the Early Cretaceous (Aptian–Albian) Ilek Formation at the Shestakovo locality in Western Siberia, Russia.It shows some primitive characters, such as slender metatarsals, a relatively long second pedal ungual, and three claws.In the likely presence of the laterodistal process on the first metatarsal the Shestakovo sauropod is similar with diplodocoids, but its more elongated and gracile first metatarsal resembles brachiosaurids (Brachiosaurus, Pleurocoelus, and Cedarosaurus), titanosaurids (Laplatasaurus), and Euhelopus. Pleurocoelus−like isolated teeth from the Shestakovo assemblage may support the brachiosaurid affinities of the Shestakovo sauropod, but a strongly procoelous mid−caudal vertebra from another locality in the same formation establishes the presence of a titanosaurid in the fauna.The foot described is referred here to as Titanosauriformes gen.et sp.indet.
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Earliest laurasian sauropod eggshells

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Megaloolithid eggshells, known from many Cretaceous deposits since 19th century, are now recognized as remnants of sauropod dinosaurs. Our paper reports the discovery of megaloolithid egg remains from the Middle Jurassic (Bajocian) of the Quercy area (southwestern France). The new Jurassic ootaxon differs from related Cretaceous oospecies in having unusually thin shells. Even Megaloolithus aureliensis, the thinnest Cretaceous megaloolithid from France is three times thicker than the Jurassic eggshells. The cladistic analysis of ootaxa reveals a peculiar point in contradiction with the phylogenetic results based on skeletal remains: the Megaloolithidae belonged to sauropod dinosaurs, which appear to be the sister group of the hadrosaur eggs (Spheroolithidae oofamilly). This result could indicate a significant amount of homoplasy in the evolution of eggshell structures, depending strongly on the incubation environment (particularly for some characters as ornamentation, pore openings and pore canals), the reproductive physiology and the oviduct function. The Bajocian eggshells might represent the earliest offshoot of the Megaloolithidae oofamily and represent the earliest sauropod eggshell record known from the deposits of Laurasia supercontinent.
The first unambiguous evidence of the presence of osteoderms in sauropod dinosaurs came from the discovery of Saltasaurus loricatus, a titanosaur from the Upper Cretaceous of Argentina. The dermal armor of Saltasaurus is composed of bony plates and small dermal ossicles. Here, we analyze the bone microstructure of these elements and provide information regarding its origin and development. The bony plates are composed almost entirely of reconstructed cancellous bone. Remains of primary bone consist of coarse bundles of mineralized collagenous fibers towards the external surface. Also, woven fibered bone tissue appears in the basal and lateral regions. Dermal ossicles lack secondary remodeling, and their matrix is formed by three orthogonal systems of collagenous fiber bundles. Growth lines are present in both bony plates and ossicles. Bone histology reveals that osteoderms mainly originated through direct mineralization (metaplasia) of the dermis, although other mechanisms are also involved (at least in the origin of dermal plates). The common features of development and integumental location of the osteoderms of Saltasaurus and other non−related vertebrates (e.g., lepidosaurs, crocodylomorphs) are linked to the intrinsic skeletogenic properties of the dermis.
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.
The Late Cretaceous South American sauropods Neuquensaurus australis and Saltasaurus loricatus are represented by well−preserved and abundant material that has been integral to our understanding of titanosaur anatomy for decades. Although the hypodigms for these species span most of the skeleton, holotypic materials are limited to a few bones that do not overlap between the two taxa. In this contribution, we augment the holotype of Neuquensaurus australis with a partial sacrum that was preserved in articulation with one of the caudal vertebrae from its original description, but not recognised as such at the time. We document this field association via the presence of a broken piece of matrix on the sixth sacral vertebral centrum that has a snap−fit to matrix on the rim of the anterior condyle of the holotypic biconvex vertebra. Based on comparisons with a more complete sacrum and ilium of a referred specimen of Neuquensaurus australis, we interpret this biconvex vertebra to be the seventh sacral vertebra. This raises the possibility that the biconvex “first caudal” vertebra of some other titanosaurs may be part of the sacrum as well. Augmentation of the Neuquensaurus australis holotype to include a sacrum makes it directly comparable to the holotype of Saltasaurus loricatus. Morphological differences in the number, shape, and proportion of sacral vertebrae allow discrimination between Neuquensaurus and Saltasaurus, confirming their generic separation. The El Brete quarry, which preserves the holotypic sacrum and abundant referred specimens of Saltasaurus loricatus, also preserves a sacrum consisting of seven vertebrae that bears autapomorphies of Neuquensaurus australis, indicating that these two saltasaurines coexisted.
Brontomerus mcintoshi is a new genus and species of sauropod dinosaur from the Hotel Mesa Quarry in Grand County, Utah, USA, in the upper part of the Ruby Ranch Member (Aptian–Albian) of the Lower Cretaceous Cedar Mountain Formation. It is known from at least two fragmentary specimens of different sizes. The type specimen is OMNH 66430, the left ilium of a juvenile individual; tentatively referred specimens include a crushed presacral centrum, a complete and well−preserved mid−to−posterior caudal vertebra, the partial centrum of a distal caudal vertebra, a complete pneumatic anterior dorsal rib from the right side, the nearly complete left scapula of a much larger, presumably adult, individual, and two partial sternal plates. Brontomerus is diagnosed by five autapomorphies of the type specimen: preacetabular lobe 55% of total ilium length, longer than in any other sauropod; preacetabular lobe directed anterolaterally at 30 to the sagittal, but straight in dorsal view and vertically oriented; postacetabular lobe reduced to near absence; ischiadic peduncle reduced to very low bulge; ilium proportionally taller than in any other sauropod, 52% as high as long. In a phylogenetic analysis, Brontomerus was recovered as a camarasauromorph in all most parsimonious trees, but with uncertain position within that clade. The large preacetabular lobe of the ilium anchored powerful protractor and abductor muscles, but precise interpretation is impossible without functionally related elements such as femora and proximal caudal vertebrae. Brontomerus is the eighth sauropod genus named from the Early Cretaceous of North America, and more remain to be described: North American sauropod diversity did not decline catastrophically at the end of the Jurassic as often assumed. The most striking differences between Late Jurassic and Early Cretaceous sauropod faunas in North America is that the former are abundant and dominated by diplodocids, whereas the latter are comparatively scarce— though still diverse—and dominated by macronarians.
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.
Many sauropod ghost lineages cross the Middle Jurassic, indicating a time interval that requires increased sampling. A wide taxonomic spectrum of sauropodomorphs is known from the Middle Jurassic of China, but the braincase of a new sauropod, named here Nebulasaurus taito gen. et sp. nov., is distinct. Nebulasaurus is sister taxon to Spinophorosaurus from the Middle Jurassic of Africa and represents a clade of basal eusauropods previously unknown from Asia. The revised faunal list indicates dramatic transitions in sauropodomorph faunas from the Jurassic to Cretaceous of Asia; these are consistent with geographic isolation of Asia through the Late Jurassic. Non-sauropod sauropodomorphs, non-mamenchisaurid eusauropods (including basal macronarians), and mamenchisaurids successively replaced previous grades through the Jurassic, and titanosauriforms excluded all other sauropod lineages across the Jurassic–Cretaceous boundary.
A new sauropod titanosaur from the Upper Cretaceous Anacleto Formation is described. Narambuenatitan palomoi gen. et sp. nov., is diagnosed by cranial and axial autapomorphies. The holotype, which represent a subadult individual, consists of the left premaxilla and maxilla, braincase, both quadrates, one cervical vertebrae, one dorsal vertebra, fragments of cervical and dorsal ribs, seventeen caudal vertebrae, caudal transverse processes, fragments of haemal arches, left sternal plate, right coracoid, left humerus, left ulnae, both pubes, iliac pedicel, proximal fragment of right ischia, and an incomplete left femur. The phylogenetic analysis indicates that Narambuenatitan is a non−eutitanosaurian lithostrotian, and that it shares with Epachthosaurus a neural spine in middle caudal vertebrae which are laminar and posteriorly elongated.
Several types of pathological bony overgrowth are known from various dinosaur taxa but, except for stress fractures, are rarely reported from appendicular elements. Herein we describe pathological manual and pedal phalanges of a camarasaurid sauropod (SMA 0002), which show features rarely recognised in non-avian dinosaurs. They include lateral osteophytes and smoothing of phalangeal articular surfaces, a deep pit, proximal enthesophytes in pedal unguals, distal overgrowth associated with a fracture, and a knob-like overgrowth lateral to the distal condyles of a pedal phalanx. Their causes were assessed by means of visual examination, CT scans, and bone histology, where possible. The lateral osteophytes are interpreted as symptoms of osteoarthritis. The ossified tendon insertions in the unguals are most probably the result of prolonged, heavy use of the pedal claws, possibly for scratch-digging. The distal overgrowth is interpreted to have developed due to changed stress regimes, and to be the cause for the fracture. The deep pit represents most likely a case of osteochondrosis, whereas the knob-like overgrowth likely represents a post-traumatic phenomenon not previously reported in dinosaurs. The study confirms that a rigorous assessment of pathologies can yield information about behaviour in long-extinct animals.
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.
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