Sauropoda
Sauropods | |
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Mounted skeleton of Apatosaurus louisae, Carnegie Museum
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | †Sauropodomorpha |
Clade: | †Anchisauria |
Clade: | †Sauropoda Marsh, 1878 |
Subgroups[1] | |
Synonyms | |
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Sauropoda (/sɔːˈrɒpədə/), whose members are known as sauropods (/ˈsɔːrəpɒdz/;[2][3] from sauro- + -pod, 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their body), 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 Apatosaurus, Argentinosaurus, Alamosaurus, Brachiosaurus, Camarasaurus, Diplodocus, and Mamenchisaurus.[4][5]
The oldest known unequivocal sauropod dinosaurs are known from the
The name Sauropoda was coined by Othniel Charles Marsh in 1878, and is derived from Ancient Greek, meaning "lizard foot".[15] Sauropods are one of the most recognizable groups of dinosaurs, and have become a fixture in popular culture due to their impressive size.
Complete sauropod fossil finds are extremely rare. Many species, especially the largest, are known only from isolated and disarticulated bones. Many near-complete specimens lack heads, tail tips and limbs.
Description
Sauropods were
Size
The sauropods' most defining characteristic was their size. Even the dwarf sauropods (perhaps 5 to 6 metres, or 20 feet 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
The longest terrestrial animal alive today, the African elephant, can only reach lengths of 7.3 metres (24 ft).[27]
Others, like the brachiosaurids, were extremely tall, with high shoulders and extremely long necks. The tallest sauropod was the giant Barosaurus specimen at 22 m (72 ft) tall.[24] By comparison, the giraffe, the tallest of all living land animals, is only 4.8 to 5.6 metres (15.74 to 18.3 ft) tall.
The best evidence indicates that the most massive were
Among the smallest sauropods were the primitive
Fossils from perhaps the largest dinosaur ever found were discovered in 2012 in the Neuquén Province of northwest Patagonia, Argentina. It is believed that they are from a titanosaur, which were amongst the largest sauropods.[36][31]
On or shortly before 29 March 2017 a sauropod footprint about 5.6 feet (1.7 meters) long was found at
Limbs and feet
As massive
The arrangement of the forefoot bone (
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.[41]
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.[42] 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.[41]
Matthew Bonnan[43][44] 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
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.[47]
Armor
Some sauropods had
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.[48] 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.[49]
It was also noted by D'Emic and his team that the differences between the teeth of the sauropods also indicated a difference in diet. Diplodocus ate plants low to the ground and Camarasaurus browsed leaves from top and middle branches. According to the scientists, the specializing of their diets helped the different herbivorous dinosaurs to coexist.[48][49]
Necks
Sauropod necks have been found at over 15 metres (49 ft) in length, a full six times longer than the world record giraffe neck.[46] 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.[46] According to Kent Stevens, computer-modeled reconstructions of the skeletons made from the vertebrae indicate that sauropod necks were capable of sweeping out large feeding areas without needing to move their bodies, but were unable to be retracted to a position much above the shoulders for exploring the area or reaching higher.[50]
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.[51] 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.[52]
Palaeobiology
Ecology
When sauropods were first discovered, their immense size led many scientists to compare them with modern-day
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.[53]
Evidence for swimming in sauropods comes from fossil trackways that have occasionally been found to preserve only the forefeet (manus) impressions. Henderson showed that such trackways can be explained by sauropods with long forelimbs (such as macronarians) floating in relatively shallow water deep enough to keep the shorter hind legs free of the bottom, and using the front limbs to punt forward.[53] However, due to their body proportions, floating sauropods would also have been very unstable and maladapted for extended periods in the water. This mode of aquatic locomotion, combined with its instability, led Henderson to refer to sauropods in water as "tipsy punters".[53]
While sauropods could therefore not have been aquatic as historically depicted, there is evidence that they preferred wet and coastal habitats. Sauropod footprints are commonly found following coastlines or crossing floodplains, and sauropod fossils are often found in wet environments or intermingled with fossils of marine organisms.
Herding and parental care
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.[58]
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.[58]
Since the segregation of juveniles and adults must have taken place soon after hatching, and combined with the fact that sauropod hatchlings were most likely
Rearing stance
Since early in the history of their study, scientists, such as
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
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,[63] such as feeding-related activities (contra Rothschild and Molnar).[62]
Head and neck posture
There is little agreement over how sauropods held their heads and necks, and the postures they could achieve in life.
Whether sauropods' long necks could be used for browsing high trees has been questioned based on calculations suggesting that just pumping blood up to the head in such a posture[64] for long would have used some half of its energy intake.[65] Further, to move blood to such a height—dismissing posited auxiliary hearts in the neck[66]—would require a heart 15 times as large as of a similar-sized whale.[67]
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".[68]
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"[68] is deeply unreliable.[69][70] Meanwhile, computer modeling of ostrich necks has raised doubts over the flexibility needed for stationary grazing.[71][72][73]
Trackways and locomotion
Sauropod
Sauropod tracks from the
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.[77] 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.[78] 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
In a study published in PLoS ONE on October 30, 2013, by
To estimate the gait and speed of Argentinosaurus, the study performed a musculoskeletal analysis. The only previous musculoskeletal analyses were conducted on
Body size
Sauropods were gigantic descendants of surprisingly small ancestors. Basal dinosauriformes, such as
Evolving from sauropodomorphs, the sauropods were huge. Their giant size probably resulted from an increased growth rate made possible by
Size in Neosauropoda
Independent gigantism
Although in general, sauropods were large, a gigantic size (40 t (39 long tons; 44 short tons) or more) was reached independently at multiple times in their evolution. Many gigantic forms existed in the Late Jurassic (specifically Kimmeridgian), such as the turiasaur Turiasaurus, the mamenchisaurids Mamenchisaurus and Xinjiangtitan, the diplodocoids Maraapunisaurus, Diplodocus, Apatosaurus, Supersaurus and Barosaurus, the camarasaurid Camarasaurus, and the brachiosaurids Brachiosaurus and Giraffatitan. Through the Early to Late Cretaceous, the giants Borealosaurus, Sauroposeidon, Paralititan, Argentinosaurus, Puertasaurus, Antarctosaurus, Dreadnoughtus, Notocolossus, Futalognkosaurus, Patagotitan and Alamosaurus lived, with all possibly being titanosaurs. One sparsely known possible giant is Huanghetitan ruyangensis, only known from 3 m (9.8 ft) long ribs. These giant species lived in the Late Jurassic to the Late Cretaceous, appearing independently over a time span of 85 million years.[51]
Dwarfism in sauropods
Two well-known island dwarf species of sauropods are the Cretaceous Magyarosaurus (at one point its identity as a dwarf was challenged) and the Jurassic Europasaurus, both from Europe. Even though these sauropods are small, the only way to prove they are true dwarfs is through a study of their bone histology. A study by Martin Sander and colleagues in 2006 examined eleven individuals of Europasaurus holgeri using bone histology and demonstrated that the small island species evolved through a decrease in the growth rate of long bones as compared to rates of growth in ancestral species on the mainland.[81] 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.[51] One of the most extreme cases of island dwarfism is found in Europasaurus, a relative of the much larger Camarasaurus and Brachiosaurus: it was only about 6.2 m (20 ft) long, an identifying trait of the species. As for all dwarf species, their reduced growth rate led to their small size.[33][51] Another taxon of tiny sauropods, the saltasaurid titanosaur Ibirania, 5.7 m (18.7 ft) long, lived a non-insular context in Upper Creaceous Brazil, and is an example of nanism resultant from other ecological pressures.[82]
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.[83][84] Sauropod ribs from Yunyang County, Chongqing, in southwest China show evidence of rib breakage by way of traumatic fracture, bone infection, and osteosclerosis.[85]
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).[86] 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.[87][88][89]
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.
The first sauropod fossil to be scientifically described was a single tooth known by the non-
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.[47]
The next sauropod find to be described and misidentified as something other than a dinosaur were a set of hip vertebrae described by Harry Seeley in 1870. Seeley found that the vertebrae were very lightly constructed for their size and contained openings for air sacs (pneumatization). Such air sacs were at the time known only in birds and pterosaurs, and Seeley considered the vertebrae to come from a pterosaur. He named the new genus Ornithopsis, or "bird face" because of this.[47]
When more complete specimens of Cetiosaurus were described by Phillips in 1871, he finally recognized the animal as a dinosaur related to Pelorosaurus.[93] 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.[94] Also in 1877, Richard Lydekker named another relative of Cetiosaurus, Titanosaurus, based on an isolated vertebra.[47]
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".[47]
Classification
The first
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.[51]
†Sauropoda |
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See also
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See also
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