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Size is an important aspect of dinosaur paleontology, of interest to both the general public and professional scientists. Dinosaurs show some of the most extreme variations in size of any land animal group, ranging from tiny hummingbirds, which can weigh as little as two grams, to the extinct titanosaurs, which could weigh as much as 50–100 t (55–110 short tons).
The latest evidence suggests that dinosaurs' average size varied through the Triassic, early Jurassic, late Jurassic and Cretaceous periods, and dinosaurs probably only became widespread during the early or mid Jurassic. [1] Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic, most often fall into the 100–1,000 kg (220–2,200 lb) category when sorted by estimated weight into categories based on order of magnitude, whereas recent predatory carnivoran mammals peak in the range of 10–100 kg (22–220 lb). [2] The mode of Mesozoic dinosaur body masses is between one and ten metric tonnes. [3] This contrasts sharply with the size of Cenozoic mammals, estimated by the National Museum of Natural History as about 2 to 5 kg (4.4 to 11.0 lb). [4]
Scientists will probably never be certain of the largest and smallest dinosaurs. This is because only a small fraction of animals ever fossilize, and most of these remains will either never be uncovered, or will be unintentionally destroyed as a result of human activity. Of the specimens that are recovered, few are even relatively complete skeletons, and impressions of skin and other soft tissues are rarely discovered. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art (though governed by some established allometric trends), and reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork, and never perfect. [5] Mass estimates for dinosaurs are much more variable than length estimates given the lack of soft tissue preservation in the fossilization process. Modern mass estimation is often done with the laser scan skeleton technique that puts a "virtual" skin over the known or implied skeleton, but the limitations inherent in previous mass estimation techniques remain. [6]
Sauropodomorph size is difficult to estimate given their usually fragmentary state of preservation. Sauropods are often preserved without their tails, so the margin of error in overall length estimates is high. Mass is calculated using the cube of the length, so for species in which the length is particularly uncertain, the weight is even more so. Estimates that are particularly uncertain (due to very fragmentary or lost material) are preceded by a question mark. Each number represents the highest estimate of a given research paper. One large sauropod, Maraapunisaurus fragillimus , was based on particularly scant remains that have been lost since their description by paleontologists in 1878. Analysis of the illustrations included in the original report suggested that M. fragillimus may have been the largest land animal of all time, possibly weighing 100–150 t (110–170 short tons) and measuring between 40–60 m (130–200 ft) long. [7] [8] One later analysis of the surviving evidence, and the biological plausibility of such a large land animal, suggested that the enormous size of this animal was an over-estimate due partly to typographical errors in the original report. [9] This would later be challenged by a different study, which argued Cope's measurements were genuine and that there was no basis for assuming typographical errors. The study, however, also reclassified the species and correspondingly gave a much lower length estimate of 30.3 metres (99 ft) and a mass of 78.5 t (86.5 short tons). [10] This in itself would later be disputed as being too small for an animal of such size, with some believing it to be even larger at around 35–40 metres (115–131 ft) and weighing around 80–120 t (88–132 short tons). [11]
Another large but even more controversial sauropod is Bruhathkayosaurus which had a calculated weight ranging between 126–220 t (139–243 short tons) and a length of 44.1 m (145 ft) [12] [13] [14] Although the existence of this sauropod had long been dismissed as a potential fake or a misidentification of a petrified tree trunk, recent photographic evidence emerged, confirming its existence. [15] More recent and reliable estimates in 2023 have rescaled Bruhathkayosaurus to weigh around 110–130 t (120–140 short tons) with its most liberal estimate being 240 t (260 short tons), making it incredibly massive for such an animal. [16] If the upper unlikely size estimates were to be taken at face value, Bruhathkayosaurus would not only be the largest dinosaur to have ever lived, but also the largest animal to have lived, exceeding even the largest blue whale recorded. According to Gregory S. Paul, 'super-sauropods' or 'land-whales' such as Maraapunisaurus, Bruhathkayosaurus and the Broome Titanosaur footprints as he calls them, should not be surprising as sauropods were more heat tolerant and grew rapidly, which allowed them to reach truly titanic sizes that rivalled the largest whales in mass despite the prevalence of air sacs. [16] Other potential factors for such extreme sauropod sizes include increasing bone robustness and load-distributing cartilaginous features to better redistribute and support such massive weights. [16]
Generally, the giant sauropods can be divided into two categories: the shorter but stockier and more massive forms (mainly titanosaurs and some brachiosaurids), and the longer but slenderer and more light-weight forms (mainly diplodocids).
Because different methods of estimation sometimes give conflicting results, mass estimates for sauropods can vary widely causing disagreement among scientists over the accurate number. For example, the titanosaur Dreadnoughtus was originally estimated to weigh 59.3 tonnes by the allometric scaling of limb-bone proportions, whereas more recent estimates, based on three-dimensional reconstructions, yield a much smaller figure of 22.1–38.2 tonnes. [17]
The sauropods were the longest and heaviest dinosaurs. For much of the dinosaur era, the smallest sauropods were larger than almost anything else in their habitat, and the largest were an order of magnitude more massive than anything else known to have walked the Earth since. Giant prehistoric mammals such as Paraceratherium and Palaeoloxodon (the largest land mammals ever discovered [18] ) were dwarfed by the giant sauropods, and only modern whales approach or surpass them in weight, though they live in the oceans. [19] There are several proposed advantages for the large size of sauropods, including protection from predation, reduction of energy use, and longevity, but it may be that the most important advantage was dietary. Large animals are more efficient at digestion than small animals, because food spends more time in their digestive systems. This also permits them to subsist on food with lower nutritive value than smaller animals. Sauropod remains are mostly found in rock formations interpreted as dry or seasonally dry, and the ability to eat large quantities of low-nutrient browse would have been advantageous in such environments. [20]
One of the tallest and heaviest dinosaurs known from good skeletons is Giraffatitan brancai (previously classified as a species of Brachiosaurus ). Its remains were discovered in Tanzania between 1907 and 1912. Bones from several similar-sized individuals were incorporated into the skeleton now mounted and on display at the Museum für Naturkunde Berlin; [21] this mount is 12–13.27 metres (39.4–43.5 ft) tall and 21.8–22.5 metres (72–74 ft) long, [22] [23] [24] and would have belonged to an animal that weighed between 30,000 to 60,000 kilograms (66,000 to 132,000 lb). One of the longest complete dinosaurs is the 27-metre-long (89 ft) Diplodocus , which was discovered in Wyoming in the United States and displayed in Pittsburgh's Carnegie Natural History Museum in 1907. [25]
There were larger dinosaurs, but knowledge of them is based entirely on a small number of fragmentary fossils. Most of the largest herbivorous specimens on record were discovered in the 1970s or later, and include the massive titanosaur Argentinosaurus huinculensis , which is the largest dinosaur known from uncontroversial and relatively substantial evidence, estimated to have been 70–80 t (77–88 short tons) and 36 m (118 ft) long. [26] [7] Some of the longest sauropods were those with exceptionally long, whip-like tails, such as the 29–30 m (95–98 ft) Diplodocus hallorum [7] [26] (formerly Seismosaurus) and the 39 m Supersaurus. [27]
In 2014, the fossilized remains of a previously unknown species of sauropod were discovered in Argentina. [28] The titanosaur, named Patagotitan mayorum , was estimated to have been around 40 m (130 ft) long weighing around 77 t (85 short tons), larger than any other previously found sauropod. The specimens found were remarkably complete, significantly more so than previous titanosaurs. It since been suggested that Patagotitan was not necessarily larger than Argentinosaurus and Puertasaurus. [29] In 2019, Patagotitan was estimated to have been 31 metres (102 ft) long and about 55 tonnes (121,000 lb) massive. [30]
The largest of non-sauropod sauropodomorphs was the unnamed 16 metres (52 ft) long 10 tonnes (22,000 lb) unnamed Elliot giant. [26] Another large sauropodomorph was Euskelosaurus . It reached 12.2 m (40 ft) in length and 2 t (2.2 short tons) in weight. [31] Yunnanosaurus youngi also reached a length of 13 m (43 ft). [32]
Tyrannosaurus was for many decades the largest and best-known theropod to the general public. Since its discovery, however, a number of other giant carnivorous dinosaurs have been described, including Spinosaurus , Carcharodontosaurus , and Giganotosaurus . [33] These large theropod dinosaurs are estimated to rival or even exceeded Tyrannosaurus rex in size, though more recent studies and reconstructions show that Tyrannosaurus, although shorter, was the bulkier animal overall. Specimens such as Sue and Scotty are both estimated to be the most massive theropods known to science. There is still no clear explanation for exactly why these animals grew so bulky and heavy compared to the land predators that came before and after them.
The largest extant theropod is the common ostrich, up to 2.74 metres (9 ft 0 in) tall and weighs between 63.5 and 145.15 kilograms (140.0 and 320.0 lb). [34]
The smallest non-avialan theropod known from adult specimens may be Anchiornis huxleyi , at 110 grams (3.9 ounces) in weight and 34 centimetres (13 in) in length, [35] although later study discovered larger specimen reaching 62 centimetres (24 in). [36] However, some studies suggest that Anchiornis was actually an avialan. [37] The smallest dinosaur known from adult specimens which is definitely not an avialan is Parvicursor remotus , at 162 grams (5.7 oz) and measuring 39 centimetres (15 in) long. [38] However, in 2022 its holotype was recognized as a juvenile individual. [39] Among living dinosaurs, the bee hummingbird (Mellisuga helenae) is smallest at 1.9 g (0.067 oz) and 5.5 cm (2.2 in) long. [40] The smallest theropod overall (including avians) is the currently extant Bee Hummingbird at 6.12 cm long and 2.6g for females, and 5.51 cm long and 3.25g for the males. [41]
In the theropod lineage leading to birds, body size shrank continuously over a period of 50 million years, from an average of 163 kilograms (359 lb) down to 0.8 kg (1.8 lb). This was the only dinosaur lineage to get continuously smaller over such an extended time period, and their skeletons developed adaptations at about four times the average rate for dinosaurs. [42] [43]
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is a subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 mya and their dominance continued throughout the Jurassic and Cretaceous periods. The fossil record shows that birds are feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch, and are the only dinosaur lineage known to have survived the Cretaceous–Paleogene extinction event approximately 66 mya. Dinosaurs can therefore be divided into avian dinosaurs—birds—and the extinct non-avian dinosaurs, which are all dinosaurs other than birds.
Argentinosaurus is a genus of giant sauropod dinosaur that lived during the Late Cretaceous period in what is now Argentina. Although it is only known from fragmentary remains, Argentinosaurus is one of the largest known land animals of all time, perhaps the largest, measuring 30–35 metres (98–115 ft) long and weighing 65–80 tonnes. It was a member of Titanosauria, the dominant group of sauropods during the Cretaceous. It is widely regarded by many paleontologists as the biggest dinosaur ever, and perhaps lengthwise the longest animal ever, though both claims have no concrete evidence yet.
Sauropoda, whose members are known as 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 Apatosaurus, Argentinosaurus, Alamosaurus, Brachiosaurus, Camarasaurus, Diplodocus, and Mamenchisaurus.
Giganotosaurus is a genus of theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period, approximately 99.6 to 95 million years ago. The holotype specimen was discovered in the Candeleros Formation of Patagonia in 1993 and is almost 70% complete. The animal was named Giganotosaurus carolinii in 1995; the genus name translates to "giant southern lizard", and the specific name honors the discoverer, Ruben Carolini. A dentary bone, a tooth, and some tracks, discovered before the holotype, were later assigned to this animal. The genus attracted much interest and became part of a scientific debate about the maximum sizes of theropod dinosaurs.
Sauroposeidon is a genus of sauropod dinosaur known from several incomplete specimens including a bone bed and fossilized trackways that have been found in the U.S. states of Oklahoma, Wyoming, and Texas.
Bruhathkayosaurus is a controversial genus of sauropod dinosaur found in the Kallamedu Formation of India. The fragmentary remains were originally described as a theropod, but it was later determined to be a titanosaurian sauropod. Length estimates by researchers exceed those of the titanosaur Argentinosaurus, as longer than 35 metres (115 ft) and weighing over 80 tonnes. A 2023 estimate placed Bruhathkayosaurus as potentially weighing approximately 110–170 tonnes. If the upper estimates of the 2023 records are accurate, Bruhathkayosaurus may have rivalled the blue whale as one of the largest animals to ever exist. However, all of the estimates are based on the dimensions of the fossils described in Yadagiri and Ayyasami (1987), and in 2017, it was reported that the holotype fossils had disintegrated and no longer exist.
Rapetosaurus is a genus of titanosaurian sauropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period. Only one species, Rapetosaurus krausei, has been identified.
Mussaurus is a genus of herbivorous sauropodomorph dinosaur that lived in southern Argentina during the Sinemurian stage of the Early Jurassic. It receives its name from the small size of the skeletons of juvenile and infant individuals, which were once the only known specimens of the genus. However, since Mussaurus is now known from adult specimens, the name is something of a misnomer.
Antarctosaurus is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now South America. The type species, Antarctosaurus wichmannianus, and a second species, Antarctosaurus giganteus, were described by prolific German paleontologist Friedrich von Huene in 1929. Three additional species of Antarctosaurus have been named since then but later studies have considered them dubious or unlikely to pertain to the genus.
Alamosaurus is a genus of opisthocoelicaudiine titanosaurian sauropod dinosaurs containing a single known species, Alamosaurus sanjuanensis, from the Maastrichtian age of the Late Cretaceous period in what is now southwestern North America. Isolated vertebrae and limb bones indicate that it reached sizes comparable to Argentinosaurus and Puertasaurus, which would make it the absolute largest dinosaur known from North America. Its fossils have been recovered from a variety of rock formations spanning the Maastrichtian age. Specimens of a juvenile Alamosaurus sanjuanensis have been recovered from only a few meters below the Cretaceous-Paleogene boundary in Texas, making it among the last surviving non-avian dinosaur species. Alamosaurus is the only known sauropod to have inhabited North America after their nearly 30-million year absence from the North American fossil record and probably represents an immigrant from South America.
Opisthocoelicaudia is a genus of sauropod dinosaur of the Late Cretaceous Period discovered in the Gobi Desert of Mongolia. The type species is Opisthocoelicaudia skarzynskii. A well-preserved skeleton lacking only the head and neck was unearthed in 1965 by Polish and Mongolian scientists, making Opisthocoelicaudia one of the best known sauropods from the Late Cretaceous. Tooth marks on this skeleton indicate that large carnivorous dinosaurs had fed on the carcass and possibly had carried away the now-missing parts. To date, only two additional, much less complete specimens are known, including part of a shoulder and a fragmentary tail. A relatively small sauropod, Opisthocoelicaudia measured about 11.4–13 m (37–43 ft) in length. Like other sauropods, it would have been characterised by a small head sitting on a very long neck and a barrel shaped trunk carried by four column-like legs. The name Opisthocoelicaudia means "posterior cavity tail", alluding to the unusual, opisthocoel condition of the anterior tail vertebrae that were concave on their posterior sides. This and other skeletal features lead researchers to propose that Opisthocoelicaudia was able to rear on its hindlegs.
Gobititan is a genus of herbivorous sauropod dinosaur from the Aptian faunal stage of the Early Cretaceous. The name of this genus is derived from the Gobi desert region and the Titans of Greek mythology, which is a reference to its large body size. The specific name shenzhouensis, is derived from "Shenzhou", an ancient name for China.
Puertasaurus is a genus of sauropod dinosaur that lived in South America during the Late Cretaceous Period. It is known from a single specimen recovered from sedimentary rocks of the Cerro Fortaleza Formation in southwestern Patagonia, Argentina, which probably is Campanian or Maastrichtian in age. The only species is Puertasaurus reuili. Described by the paleontologist Fernando Novas and colleagues in 2005, it was named in honor of Pablo Puerta and Santiago Reuil, who discovered and prepared the specimen. It consists of four well-preserved vertebrae, including one cervical, one dorsal, and two caudal vertebrae. Puertasaurus is a member of Titanosauria, the dominant group of sauropods during the Cretaceous.
Europasaurus is a basal macronarian sauropod, a form of quadrupedal herbivorous dinosaur. It lived during the Late Jurassic of northern Germany, and has been identified as an example of insular dwarfism resulting from the isolation of a sauropod population on an island within the Lower Saxony basin.
Nigersaurus is a genus of rebbachisaurid sauropod dinosaur that lived during the middle Cretaceous period, about 115 to 105 million years ago. It was discovered in the Elrhaz Formation in an area called Gadoufaoua, in Niger. Fossils of this dinosaur were first described in 1976, but it was only named Nigersaurus taqueti in 1999, after further and more complete remains were found and described. The genus name means "Niger reptile", and the specific name honours the palaeontologist Philippe Taquet, who discovered the first remains.
Paralititan was a giant titanosaurian sauropod dinosaur genus discovered in coastal deposits in the Upper Cretaceous Bahariya Formation of Egypt. It lived between 99.6 and 93.5 million years ago.
Patagotitan is a genus of titanosaurian sauropod dinosaur from the Cerro Barcino Formation in Chubut Province, Patagonia, Argentina. The genus contains a single species known from at least six young adult individuals, Patagotitan mayorum, which was first announced in 2014 and then named in 2017 by José Carballido and colleagues. Preliminary studies and press releases suggested that Patagotitan was the largest known titanosaur and land animal overall, with an estimated length of 37 m (121 ft) and an estimated weight of 69 tonnes. Later research revised the length estimate down to 31 m (102 ft) and weight estimates down to approximately 50–57 tonnes, suggesting that Patagotitan was of a similar size to, if not smaller than, its closest relatives Argentinosaurus and Puertasaurus. Still, Patagotitan is one of the most-known titanosaurs, and so its interrelationships with other titanosaurs have been relatively consistent in phylogenetic analyses. This led to its use in a re-definition of the group Colossosauria by Carballido and colleagues in 2022.
Dreadnoughtus is a genus of titanosaurian sauropod dinosaur containing a single species, Dreadnoughtus schrani. D. schrani is known from two partial skeletons discovered in Upper Cretaceous rocks of the Cerro Fortaleza Formation in Santa Cruz Province, Argentina. It is one of the largest terrestrial vertebrates known, with the immature type specimen measuring 26 metres (85 ft) in total body length and weighing 48–49 metric tons. D. schrani is known from more complete skeletons than any other gigantic titanosaurian.
Notocolossus is a genus of titanosaurian sauropod dinosaur from late Cretaceous strata of Mendoza Province, Argentina.
Maraapunisaurus is a genus of sauropod dinosaur from the Late Jurassic Morrison Formation of western North America. Originally named Amphicoelias fragillimus, it has sometimes been estimated to be the largest dinosaur specimen ever discovered. Based on surviving descriptions of a single fossil bone, scientists have produced numerous size estimates over the years; the largest estimate M. fragillimus to have been the longest known animal at 58 metres (190 ft) in length with a mass of 150 tonnes. However, because the only fossil remains were lost at some point after being studied and described in the 1870s, evidence survived only in contemporary drawings and field notes.
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