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Sauropoda

Sauropoda, whose members are called sauropods, is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads, and four thick, pillar-like legs. They are notable for the enormous sizes attained by some species, and the group includes the largest animals to have ever lived on land. Well-known genera include Alamosaurus, Apatosaurus, Argentinosaurus, Brachiosaurus, Brontosaurus, Camarasaurus, Diplodocus, Dreadnoughtus, and Mamenchisaurus.

Description
Sauropods were herbivorous (plant-eating), usually quite long-necked quadrupeds (four-legged), often with spatulate (spatula-shaped: broad at the tip, narrow at the neck) teeth. They had relatively tiny heads, massive bodies, and most had long tails. Their hind legs were thick, straight, and powerful, ending in club-like feet with five toes, though only the inner three (or in some cases four) bore claws. Their forelimbs were rather more slender and typically ended in pillar-like hands built for supporting weight; often only the thumb bore a claw. Many illustrations of sauropods in the flesh miss these facts, inaccurately depicting sauropods with hooves capping the claw-less digits of the feet, or more than three claws or hooves on the hands. The proximal caudal vertebrae are extremely diagnostic for sauropods. Size The sauropods' most defining characteristic was their size. Even the dwarf sauropods (perhaps long) were counted among the largest animals in their ecosystem. Their only real competitors in terms of size are the rorquals, such as the blue whale. But, unlike whales, sauropods were primarily terrestrial animals. Their body structure did not vary as much as other dinosaurs, perhaps due to size constraints, but they displayed ample variety. Some, like the diplodocids, possessed tremendously long tails, which they may have been able to crack like a whip as a signal or to deter or injure predators, or to make sonic booms. Supersaurus, at long, was the longest sauropod known from reasonably complete remains, but others, like the old record holder, Diplodocus, were also extremely long. The holotype (and now lost) vertebra of Amphicoelias fragillimus (now Maraapunisaurus) may have come from an animal long; its vertebral column would have been substantially longer than that of the blue whale. However, research published in 2015 speculated that the size estimates of A. fragillimus may have been highly exaggerated. The longest dinosaur known from reasonable fossils material is probably Argentinosaurus huinculensis with length estimates of according to the most recent researches. However the giant Barosaurus specimen BYU 9024 might have been even larger reaching lengths of . Others, like the brachiosaurids, were extremely tall, with high shoulders and extremely long necks. The tallest sauropod was the giant Barosaurus specimen at tall. allowing for more updated estimates that range between , rivaling the blue whale in size. The weight of Amphicoelias fragillimus was estimated at tons with lengths of up to nearly The largest land animal alive today, the bush elephant, weighs no more than . Among the smallest sauropods were the primitive Ohmdenosaurus ( long), the dwarf titanosaur Magyarosaurus ( long), and the dwarf brachiosaurid Europasaurus, which was long as a fully-grown adult. Its small stature was probably the result of insular dwarfism occurring in a population of sauropods isolated on an island of the late Jurassic in what is now the Langenberg area of northern Germany. The diplodocoid sauropod Brachytrachelopan was the shortest member of its group because of its unusually short neck. Unlike other sauropods, whose necks could grow to up to four times the length of their backs, the neck of Brachytrachelopan was shorter than its backbone. Fossils from perhaps the largest dinosaur ever found (MOZ-Pv 1221) were discovered in 2021 in the Neuquén Province of northwest Patagonia, Argentina. It is believed that they are from a titanosaur, which were amongst the largest sauropods. The report said that it was the biggest known yet. In 2020 Molina-Perez and Larramendi estimated the size of the animal at and 72 tonnes (79.4 short tons) based on the long footprint. Limbs and feet As massive quadrupeds, sauropods developed specialized "graviportal" (weight-bearing) limbs. The hind feet were broad, and retained three claws in most species. Particularly unusual compared with other animals were the highly modified front feet (manus). The front feet of sauropods were very dissimilar from those of modern large quadrupeds, such as elephants. Rather than splaying out to the sides to create a wide foot as in elephants, the manus bones of sauropods were arranged in fully vertical columns, with extremely reduced finger bones (though it is not clear if the most primitive sauropods, such as Vulcanodon and Barapasaurus, had such forefeet). The front feet were so modified in eusauropods that individual digits would not have been visible in life. The arrangement of the forefoot bone (metacarpal) columns in eusauropods was semi-circular, so sauropod forefoot prints are horseshoe-shaped. Unlike elephants, print evidence shows that sauropods lacked any fleshy padding to back the front feet, making them concave. Titanosaurs may have lost the thumb claw completely (with the exception of early forms, such as Janenschia). Titanosaurs were most unusual among sauropods, as, across their history as a clade, they lost not just the external claw but also completely lost the digits of the front foot. Advanced titanosaurs had no digits or digit bones, and walked only on horseshoe-shaped "stumps" made up of the columnar metacarpal bones. Print evidence from Portugal shows that, in at least some sauropods (probably brachiosaurids), the bottom and sides of the forefoot column was likely covered in small, spiny scales, which left score marks in the prints. In titanosaurs, the ends of the metacarpal bones that contacted the ground were unusually broad and squared-off, and some specimens preserve the remains of soft tissue covering this area, suggesting that the front feet were rimmed with some kind of padding in these species. has shown that sauropod dinosaur long bones grew isometrically: that is, there was little to no change in shape as juvenile sauropods became gigantic adults. Bonnan suggested that this odd scaling pattern (most vertebrates show significant shape changes in long bones associated with increasing weight support) might be related to a stilt-walker principle (suggested by amateur scientist Jim Schmidt) in which the long legs of adult sauropods allowed them to easily cover great distances without changing their overall mechanics. Air sacs Along with other saurischian dinosaurs (such as theropods, including birds), sauropods had a system of air sacs, evidenced by indentations and hollow cavities in most of their vertebrae that had been invaded by them. Pneumatic, hollow bones are a characteristic feature of all sauropods. These air spaces reduced the overall weight of the massive necks that the sauropods had, and the air-sac system in general, allowing for a single-direction airflow through stiff lungs, made it possible for the sauropods to get enough oxygen. The bird-like hollowing of sauropod bones was recognized early in the study of these animals, and, in fact, at least one sauropod specimen found in the 19th century (Ornithopsis) was originally misidentified as a pterosaur (a flying reptile) because of this. Armor '' Some sauropods had armor. There were genera with small clubs on their tails, a prominent example being Shunosaurus, and several titanosaurs, such as Saltasaurus and Ampelosaurus, had small bony osteoderms covering portions of their bodies. Teeth A study by Michael D'Emic and his colleagues from Stony Brook University found that sauropods evolved high tooth replacement rates to keep up with their large appetites. The study suggested that Nigersaurus, for example, replaced each tooth every 14 days, Camarasaurus replaced each tooth every 62 days, and Diplodocus replaced each tooth once every 35 days. The scientists found qualities of the tooth affected how long it took for a new tooth to grow. Camarasaurus's teeth took longer to grow than those for Diplodocus because they were larger. Enabling this were a number of essential physiological features. The dinosaurs' overall large body size and quadrupedal stance provided a stable base to support the neck, and the head was evolved to be very small and light, losing the ability to orally process food. By reducing their heads to simple harvesting tools that got the plants into the body, the sauropods needed less power to lift their heads, and thus were able to develop necks with less dense muscle and connective tissue. This drastically reduced the overall mass of the neck, enabling further elongation. Sauropods also had a great number of adaptations in their skeletal structure. Some sauropods had as many as 19 cervical vertebrae, whereas almost all mammals are limited to only seven. Additionally, each vertebra was extremely long and had a number of empty spaces in them which would have been filled only with air. An air-sac system connected to the spaces not only lightened the long necks, but effectively increased the airflow through the trachea, helping the creatures to breathe in enough air. By evolving vertebrae consisting of 60% air, the sauropods were able to minimize the amount of dense, heavy bone without sacrificing the ability to take sufficiently large breaths to fuel the entire body with oxygen. Another proposed function of the sauropods' long necks was essentially a radiator to deal with the extreme amount of heat produced from their large body mass. Considering that the metabolism would have been doing an immense amount of work, it would certainly have generated a large amount of heat as well, and elimination of this excess heat would have been essential for survival. It has also been proposed that the long necks would have cooled the veins and arteries going to the brain, avoiding excessively heated blood from reaching the head. It was in fact found that the increase in metabolic rate resulting from the sauropods' necks was slightly more than compensated for by the extra surface area from which heat could dissipate. ==Palaeobiology==
Palaeobiology
Ecology Dental microwear texture analysis (DMTA) performed on a titanosauriform sauropod from the Turonian-aged Tamagawa Formation suggests that the sauropod fed on plant material that was softer than insect exoskeletons or mollusc shells, with the diet likely consisting of ferns and gymnosperms. The DMTA results also suggested that sauropods likely masticated more energetically than present-day lepidosaurs do. When sauropods were first discovered, their immense size led many scientists to compare them with modern-day whales. Most studies in the 19th and early 20th centuries concluded that sauropods were too large to have supported their weight on land, and therefore that they must have been mainly aquatic. Most life restorations of sauropods in art through the first three quarters of the 20th century depicted them fully or partially immersed in water. This early notion was cast in doubt beginning in the 1950s, when a study by Kermack (1951) demonstrated that, if the animal were submerged in several metres of water, the pressure would be enough to fatally collapse the lungs and airway. However, this and other early studies of sauropod ecology were flawed in that they ignored a substantial body of evidence that the bodies of sauropods were heavily permeated with air sacs. In 1878, paleontologist E.D. Cope had even referred to these structures as "floats". Beginning in the 1970s, the effects of sauropod air sacs on their supposed aquatic lifestyle began to be explored. Paleontologists such as Coombs and Bakker used this, as well as evidence from sedimentology and biomechanics, to show that sauropods were primarily terrestrial animals. In 2004, D.M. Henderson noted that, due to their extensive system of air sacs, sauropods would have been buoyant and would not have been able to submerge their torsos completely below the surface of the water; in other words, they would float, and would not have been in danger of lung collapse due to water pressure when swimming. Studies published in 2021 suggest sauropods could not inhabit polar regions. This study suggests they were largely confined to tropical areas and had metabolisms that were very different to those of other dinosaurs, perhaps intermediate between mammals and reptiles. New studies published by Taia Wyenberg-henzler in 2022 suggest that sauropods in North America declined due to undetermined reasons in regards to their niches and distribution during the end of the Jurassic and into the latest Cretaceous. Why this is remains unclear, but some similarities in feeding niches between iguanodontians, hadrosauroids, and sauropods have been suggested and may have resulted in some competition. However, this cannot fully explain the full decline in distribution of sauropods, as competitive exclusion would have resulted in a much more rapid decline than what is shown in the fossil record. Moreover, it must be determined as to whether sauropod declines in North America was the result of a change in preferred flora that sauropods ate, climate, or other factors. It is also suggested in this same study that iguanodontians and hadrosauroids took advantage of recently vacated niches left by a decline in sauropod diversity during the late Jurassic and the Cretaceous in North America. Herding and parental care '', formed herds segregated by age. Many lines of fossil evidence, from both bone beds and trackways, indicate that sauropods were gregarious animals that formed herds. However, the makeup of the herds varied between species. Some bone beds, for example a site from the Middle Jurassic of Argentina, appear to show herds made up of individuals of various age groups, mixing juveniles and adults. However, a number of other fossil sites and trackways indicate that many sauropod species travelled in herds segregated by age, with juveniles forming herds separate from adults. Such segregated herding strategies have been found in species such as Alamosaurus, Bellusaurus and some diplodocids. In a review of the evidence for various herd types, Myers and Fiorillo attempted to explain why sauropods appear to have often formed segregated herds. Studies of microscopic tooth wear show that juvenile sauropods had diets that differed from their adult counterparts, so herding together would not have been as productive as herding separately, where individual herd members could forage in a coordinated way. The vast size difference between juveniles and adults may also have played a part in the different feeding and herding strategies. A 2014 study suggested that the time from laying the egg to the time of the hatching was likely to have been between 65 and 82 days. Exactly how segregated versus age-mixed herding varied across different groups of sauropods is unknown. Further examples of gregarious behavior will need to be discovered from more sauropod species to detect possible distribution patterns. Rearing stance , 1911 Since early in the history of their study, scientists, such as Osborn, have speculated that sauropods could rear up on their hind legs, using the tail as the third 'leg' of a tripod. A skeletal mount depicting the diplodocid Barosaurus lentus rearing up on its hind legs at the American Museum of Natural History is one illustration of this hypothesis. In a 2005 paper, Rothschild and Molnar reasoned that if sauropods had adopted a bipedal posture at times, there would be evidence of stress fractures in the forelimb 'hands'. However, none were found after they examined a large number of sauropod skeletons. '', depicted in a rearing tripodal stance Heinrich Mallison (in 2009) was the first to study the physical potential for various sauropods to rear into a tripodal stance. Mallison found that some characters previously linked to rearing adaptations were actually unrelated (such as the wide-set hip bones of titanosaurs) or would have hindered rearing. For example, titanosaurs had an unusually flexible backbone, which would have decreased stability in a tripodal posture and would have put more strain on the muscles. Likewise, it is unlikely that brachiosaurids could rear up onto the hind legs, as their center of gravity was much farther forward than other sauropods, which would cause such a stance to be unstable. Diplodocids, on the other hand, appear to have been well adapted for rearing up into a tripodal stance. Diplodocids had a center of mass directly over the hips, giving them greater balance on two legs. Diplodocids also had the most mobile necks of sauropods, a well-muscled pelvic girdle, and tail vertebrae with a specialised shape that would allow the tail to bear weight at the point it touched the ground. Mallison concluded that diplodocids were better adapted to rearing than elephants, which do so occasionally in the wild. He also argues that stress fractures in the wild do not occur from everyday behaviour, for long would have used some half of its energy intake. Further, to move blood to such a height—dismissing posited auxiliary hearts in the neck—would require a heart 15 times as large as of a similar-sized whale. The above have been used to argue that the long neck must instead have been held more or less horizontally, presumed to enable feeding on plants over a wide area with less need to move about, yielding a large energy saving for such a large animal. Reconstructions of the necks of Diplodocus and Apatosaurus have therefore often portrayed them in near-horizontal, so-called "neutral, undeflected posture". However, research on living animals demonstrates that almost all extant tetrapods hold the base of their necks sharply flexed when alert, showing that any inference from bones about habitual "neutral postures" Meanwhile, computer modeling of ostrich necks has raised doubts over the flexibility needed for stationary grazing. Trackways and locomotion , Italy at Serras de Aire e Candeeiros Natural Park, Portugal Sauropod trackways and other fossil footprints (known as "ichnites") are known from abundant evidence present on most continents. Ichnites have helped support other biological hypotheses about sauropods, including general fore and hind foot anatomy (see Limbs and feet above). Generally, prints from the forefeet are much smaller than the hind feet, and often crescent-shaped. Occasionally ichnites preserve traces of the claws, and help confirm which sauropod groups lost claws or even digits on their forefeet. Sauropod tracks from the Villar del Arzobispo Formation of early Berriasian age in Spain support the gregarious behaviour of the group. The tracks are possibly more similar to Sauropodichnus giganteus than any other ichnogenera, although they have been suggested to be from a basal titanosauriform. The tracks are wide-gauge, and the grouping as close to Sauropodichnus is also supported by the manus-to-pes distance, the morphology of the manus being kidney bean-shaped, and the morphology of the pes being subtriangular. It cannot be identified whether the footprints of the herd were caused by juveniles or adults, because of the lack of previous trackway individual age identification. Generally, sauropod trackways are divided into three categories based on the distance between opposite limbs: narrow gauge, medium gauge, and wide gauge. The gauge of the trackway can help determine how wide-set the limbs of various sauropods were and how this may have impacted the way they walked. Occasionally, only trackways from the forefeet are found. Falkingham et al. used computer modelling to show that this could be due to the properties of the substrate. These need to be just right to preserve tracks. Differences in hind limb and fore limb surface area, and therefore contact pressure with the substrate, may sometimes lead to only the forefeet trackways being preserved. Biomechanics and speed , Plaza Huincul, Argentina In a study published in PLoS ONE on October 30, 2013, by Bill Sellers, Rodolfo Coria, Lee Margetts et al., Argentinosaurus was digitally reconstructed to test its locomotion for the first time. Before the study, the most common way of estimating speed was through studying bone histology and ichnology. Commonly, studies about sauropod bone histology and speed focus on the postcranial skeleton, which holds many unique features, such as an enlarged process on the ulna, a wide lobe on the ilia, an inward-slanting top third of the femur, and an extremely ovoid femur shaft. Those features are useful when attempting to explain trackway patterns of graviportal animals. When studying ichnology to calculate sauropod speed, there are a few problems, such as only providing estimates for certain gaits because of preservation bias, and being subject to many more accuracy problems. To estimate the gait and speed of Argentinosaurus, the study performed a musculoskeletal analysis. The only previous musculoskeletal analyses were conducted on hominoids, terror birds, and other dinosaurs. Before they could conduct the analysis, the team had to create a digital skeleton of the animal in question, show where there would be muscle layering, locate the muscles and joints, and finally find the muscle properties before finding the gait and speed. The results of the biomechanics study revealed that Argentinosaurus was mechanically competent at a top speed of 2 m/s (5 mph) given the great weight of the animal and the strain that its joints were capable of bearing. The results further revealed that much larger terrestrial vertebrates might be possible, but would require significant body remodeling and possible sufficient behavioral change to prevent joint collapse. Two other possible dwarfs are Rapetosaurus, which existed on the island of Madagascar, an isolated island in the Cretaceous, and Ampelosaurus, a titanosaur that lived on the Iberian peninsula of southern Spain and France. Amanzia from Switzerland might also be a dwarf, but this has yet to be proven. Paleopathology and paleoparasitology Sauropods are rarely known for preserved injuries or signs of illnesses, but more recent discoveries show they could suffer from such pathologies. A diplodocid specimen from the Morrison Formation referred to as "Dolly" was described in 2022 with evidence of a severe respiratory infection. Sauropod ribs from Yunyang County, Chongqing, in southwest China show evidence of rib breakage by way of traumatic fracture, bone infection, and osteosclerosis. A sauropod tibia exhibiting initial fracture has been described from the Middle Jurassic of Yunyang County in southwestern China. Ibirania, a nanoid titanosaur fossil from Brazil, suggests that individuals of various genera were susceptible to diseases such as osteomyelitis and parasite infestations. The specimen hails from the late cretaceous São José do Rio Preto Formation, Bauru Basin, and was described in the journal Cretaceous Research by Aureliano et al. (2021). Examination of the titanosaur's bones revealed what appear to be parasitic blood worms similar to the prehistoric Paleoleishmania but are 10-100 times larger, that seemed to have caused the osteomyelitis. The fossil is the first known instance of an aggressive case of osteomyelitis being caused by blood worms in an extinct animal. ==History of discovery==
History of discovery
The first scraps of fossil remains now recognized as sauropods all came from England and were originally interpreted in a variety of different ways. Their relationship to other dinosaurs was not recognized until well after their initial discovery. ''. By John A. Ryder, 1877. The first sauropod fossil to be scientifically described was a single tooth known by the non-Linnaean descriptor Rutellum implicatum. This fossil was described by Edward Lhuyd in 1699, but was not recognized as a giant prehistoric reptile at the time. Dinosaurs would not be recognized as a group until over a century later. Richard Owen published the first modern scientific descriptions of sauropods in 1841, in a book and a paper naming Cardiodon and Cetiosaurus. Cardiodon was known only from two unusual, heart-shaped teeth (from which it got its name), which could not be identified beyond the fact that they came from a previously unknown large reptile. Cetiosaurus was known from slightly better, but still scrappy remains. Owen thought at the time that Cetiosaurus was a giant marine reptile related to modern crocodiles, hence its name, which means "whale lizard". A year later, when Owen coined the name Dinosauria, he did not include Cetiosaurus and Cardiodon in that group. In 1850, Gideon Mantell recognized the dinosaurian nature of several bones assigned to Cetiosaurus by Owen. Mantell noticed that the leg bones contained a medullary cavity, a characteristic of land animals. He assigned these specimens to the new genus Pelorosaurus, and grouped it together with the dinosaurs. However, Mantell still did not recognize the relationship to Cetiosaurus. However, it was not until the description of new, nearly complete sauropod skeletons from the United States (representing Apatosaurus and Camarasaurus) later that year that a complete picture of sauropods emerged. An approximate reconstruction of a complete sauropod skeleton was produced by artist John A. Ryder, hired by paleontologist E.D. Cope, based on the remains of Camarasaurus, though many features were still inaccurate or incomplete according to later finds and biomechanical studies. Also in 1877, Richard Lydekker named another relative of Cetiosaurus, Titanosaurus, based on an isolated vertebra. , Brachiosaurus, Giraffatitan, and Euhelopus''. In 1878, the most complete sauropod yet was found and described by Othniel Charles Marsh, who named it Diplodocus. With this find, Marsh also created a new group to contain Diplodocus, Cetiosaurus, and their increasing roster of relatives to differentiate them from the other major groups of dinosaurs. Marsh named this group Sauropoda, or "lizard feet". ==Classification==
Classification
The first phylogenetic definition of Sauropoda was published in 1997 by Salgado and colleagues. They defined the clade as a node-based taxon, containing "the most recent common ancestor of Vulcanodon karibaensis and Eusauropoda and all of its descendants". Later, several stem-based definitions were proposed, including one by Yates (2007), who defined Sauropoda as "the most inclusive clade that includes Saltasaurus loricatus but not Melanorosaurus readi". Proponents of this definition also use the clade name Gravisauria, defined as the most recent ancestor of Tazoudasaurus naimi and Saltasaurus loricatus and all of its descendants for the clade equivalent to Sauropoda as defined by Salgado et al. The clade Gravisauria was appointed by the French paleontologist Ronan Allain and Moroccan paleontologist Najat Aquesbi in 2008 when a cladistic analysis of the dinosaur found by Allain, Tazoudasaurus, as the outcome was that the family Vulcanodontidae. The group includes Tazoudasaurus and Vulcanodon, and the sister taxon Eusauropoda, but also certain species such as Antetonitrus, Gongxianosaurus and Isanosaurus that do not belong in Vulcanodontidae but to an even more basic position occupied in Sauropoda. It made sense to have Sauropoda compared to this, more derived group that included Vulcanodontidae and Eusauropoda in a definition: defined as the group formed by the last common ancestor of Tazoudasaurus and Saltasaurus (Bonaparte and Powell, 1980) and all its descendants. Aquesbi mentioned two synapomorphies, shared derived characteristics of Gravisauria: the vertebrae are wider side to side than front to rear and possession of asymmetrical condyles femoris at the bottom of the femur. Those were previously not thought to be Eusauropoda synapomorphies but Allian found these properties also on Tazoudasaurus. Gravisauria split off in the Early Jurassic, around the Pliensbachian and Toarcian, 183 million years ago, and Aquesbi thought that this was part of a much larger revolution in the fauna, which includes the disappearance of Prosauropoda, Coelophysoidea and basal Thyreophora, which they attributed to a worldwide mass extinction. The phylogenetic relationships of the sauropods have largely stabilised in recent years, though there are still some uncertainties, such as the placement of Euhelopus, Haplocanthosaurus, Jobaria and Nemegtosauridae. Cladogram after an analysis presented by Sander and colleagues in 2011. }} ==See also==
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