Brontosaurus (/ˌbrɒntəˈsɔːrəs/; meaning "thunder lizard" from Greek βροντή, brontē "thunder" and σαῦρος, sauros "lizard") is a genus of gigantic quadruped sauropod dinosaurs. Although the type species, B. excelsus, had long been considered a species of the closely related Apatosaurus, researchers proposed in 2015 that Brontosaurus is a genus separate from Apatosaurus and that it contains three species: B. excelsus, B. yahnahpin, and B. parvus.
Brontosaurus had a long, thin neck and a small head adapted for a herbivorous lifestyle, a bulky, heavy torso, and a long, whip-like tail. The various species lived during the Late Jurassic epoch, in the Morrison Formation of what is now North America, and were extinct by the end of the Jurassic. Adult individuals of Brontosaurus are estimated to have weighed up to 15 tonnes (17 short tons) and measured up to 22 metres (72 feet) long.
As the archetypal sauropod, Brontosaurus is one of the best-known dinosaurs and has been featured in film, advertising, and postage stamps, as well as many other types of media.
Description[]
Brontosaurus was a large, long-necked, quadrupedal animal with a long, whip-like tail, and fore limbs that were slightly shorter than its hind limbs. The largest species, B. excelsus, weighed up to 15 t (17 short tons) and measured up to 22 m (72 ft) long from head to tail.
The skull of Brontosaurus has not been found, but was probably similar to the skull of the closely related Apatosaurus. Like those of other sauropods, the vertebrae of the neck were deeply bifurcated; that is, they carried paired spines, resulting in a wide and deep neck. The spine and tail consisted of 15 cervicals, ten dorsals, five sacrals, and about 82 caudals. The number of caudal vertebrae was noted to vary, even within a species. The cervical vertebrae were stouter than other diplodocids, though not as stout as in mature specimens of Apatosaurus. The dorsal ribs are not fused or tightly attached to their vertebrae, instead being loosely articulated. Ten dorsal ribs are on either side of the body. The large neck was filled with an extensive system of weight-saving air sacs.[citation needed] Brontosaurus, like its close relative Apatosaurus, had tall spines on its vertebrae, which made up more than half the height of the individual bones. The shape of the tail was unusual for diplodocids, being comparatively slender, due to the vertebral spines rapidly decreasing in height the farther they are from the hips. Brontosaurus spp. also had very long ribs compared to most other diplodocids, giving them unusually deep chests. As in other diplodocids, the last portion of the tail of Brontosaurus possessed a whip-like structure.
The limb bones were also very robust. The arm bones are stout, with the humerus resembling that of Camarasaurus, and those of B. excelsus being nearly identical to those of Apatosaurus ajax. Charles Gilmore in 1936 noted that previous reconstructions erroneously proposed that the radius and ulna could cross, when in life they would have remained parallel. Brontosaurus had a single large claw on each fore limb, and the first three toes possessed claws on each foot. Even by 1936, it was recognized that no sauropod had more than one hand claw preserved, and this one claw is now accepted as the maximum number throughout the entire group. The single front claw bone is slightly curved and squarely shortened on the front end. The hip bones included robust ilia and the fused pubes and ischia. The tibia and fibula bones of the lower leg were different from the slender bones of Diplodocus, but nearly indistinguishable from those of Camarasaurus. The fibula is longer than the tibia, although it is also more slender.
History[]
In 1879, Othniel Charles Marsh, a professor of paleontology at Yale University, announced the discovery of a large and fairly complete sauropod skeleton from Morrison Formation rocks at Como Bluff, Wyoming. He identified it as belonging to an entirely new genus and species, which he named Brontosaurus excelsus, meaning "thunder lizard", from the Greek brontē/βροντη meaning "thunder" and sauros/σαυρος meaning "lizard", and from the Latin excelsus, "noble" or "high". By this time, the Morrison Formation had become the center of the Bone Wars, a fossil-collecting rivalry between Marsh and another early paleontologist, Edward Drinker Cope. Because of this, the publications and descriptions of taxa by Marsh and Cope were rushed at the time.
Elmer Riggs, in the 1903 edition of Geological Series of the Field Columbian Museum, argued that Brontosaurus was not different enough from Apatosaurus to warrant its own genus, so he created the new combination Apatosaurus excelsus for it. Riggs stated that "In view of these facts the two genera may be regarded as synonymous. As the term 'Apatosaurus' has priority, 'Brontosaurus' will be regarded as a synonym". Nonetheless, before the mounting of the American Museum of Natural History specimen, Henry Fairfield Osborn chose to label the skeleton "Brontosaurus", though he was a strong opponent of Marsh and his taxa.
In 1905, the American Museum of Natural History (AMNH) unveiled the first-ever mounted skeleton of a sauropod, a composite specimen (mainly made of bones from AMNH 460) that they referred to as the species Brontosaurus excelsus. The AMNH specimen was very complete, only missing the feet (feet from the specimen AMNH 592 were added to the mount), lower leg, and shoulder bones (added from AMNH 222), and tail bones (added from AMNH 339). To complete the mount, the rest of the tail was fashioned to appear as Marsh believed it should, which had too few vertebrae. In addition, a sculpted model of what the museum felt the skull of this massive creature might look like was placed on the skeleton. This was not a delicate skull like that of Diplodocus, which would later turn out to be more accurate, but was based on "the biggest, thickest, strongest skull bones, lower jaws and tooth crowns from three different quarries". These skulls were likely those of Camarasaurus, the only other sauropod for which good skull material was known at the time. The mount construction was overseen by Adam Hermann, who failed to find Brontosaurus skulls. Hermann was forced to sculpt a stand-in skull by hand. Henry Fairfield Osborn noted in a publication that the skull was "largely conjectural and based on that of Morosaurus" (now Camarasaurus).
In 1909, an Apatosaurus skull was found, during the first expedition to what would become the Carnegie Quarry at Dinosaur National Monument, led by Earl Douglass. The skull was found a few meters away from a skeleton (specimen CM 3018) identified as the new species Apatosaurus louisae. The skull was designated CM 11162, and was very similar to the skull of Diplodocus. It was accepted as belonging to the Apatosaurus specimen by Douglass and Carnegie Museum director William H. Holland, although other scientists, most notably Osborn, rejected this identification. Holland defended his view in 1914 in an address to the Paleontological Society of America, yet he left the Carnegie Museum mount headless. While some thought Holland was attempting to avoid conflict with Osborn, others suspected that Holland was waiting until an articulated skull and neck were found to confirm the association of the skull and skeleton. After Holland's death in 1934, a cast of a Camarasaurus skull was placed on the mount by museum staff.
At the Yale Peabody Museum, a skeleton was mounted in 1931 with a skull unique from all the others. While at the time most museums were using Camarasaurus casts, the Peabody Museum sculpted a completely different skull. They based the lower jaw on a Camarasaurus mandible, with the cranium resembling Marsh's 1891 illustration. The skull also included forward-pointing nasals, something truly different to any dinosaur, and fenestrae differing from the drawing and other skulls.
No apatosaurine skull was mentioned in literature until the 1970s, when John Stanton McIntosh and David Berman redescribed the skulls of Diplodocus and Apatosaurus. They found that though he never published his opinion, Holland was almost certainly correct, that Apatosaurus (and Brontosaurus) had a Diplodocus-like skull. According to them, many skulls long thought to pertain to Diplodocus might instead be those of Apatosaurus. They reassigned multiple skulls to Apatosaurus based on associated and closely associated vertebrae. Though they supported Holland, Apatosaurus was noted to possibly have possessed a Camarasaurus-like skull, based on a disarticulated Camarasaurus-like tooth found at the precise site where an Apatosaurus specimen was found years before. On October 20, 1979, after the publications by McIntosh and Berman, the first skull of an Apatosaurus was mounted on a skeleton in a museum, that of the Carnegie. In 1995, the American Museum of Natural History followed suit, and unveiled their remounted skeleton (now labelled Apatosaurus excelsus) with a corrected tail and a new skull cast from A. louisae. In 1998, the Felch Quarry skull that Marsh included in his 1896 skeletal restoration was suggested to belong to Brachiosaurus instead. In 2011, the first specimen of Apatosaurus where a skull was found articulated with its cervical vertebrae was described. This specimen, CMC VP 7180, was found to differ in both skull and neck features from A. louisae, and the specimen was found to have a majority of features related to those of A. ajax.
Almost all 20th-century paleontologists agreed with Riggs that all Apatosaurus and Brontosaurus species should be classified in a single genus. According to the rules of the ICZN (which governs the scientific names of animals), the name Apatosaurus, having been published first, had priority as the official name; Brontosaurus was considered a junior synonym and was therefore discarded from formal use. Despite this, at least one paleontologist—Robert T. Bakker—argued in the 1990s that A. ajax and A. excelsus are in fact sufficiently distinct that the latter continues to merit a separate genus. In 2015, an extensive study of diplodocid relationships by Emanuel Tschopp, Octavio Mateus, and Roger Benson concluded that Brontosaurus was indeed a valid genus of sauropod distinct from Apatosaurus. The scientists developed a statistical method to more objectively assess differences between fossil genera and species, and concluded that Brontosaurus could be "resurrected" as a valid name. They assigned two former Apatosaurus species, A. parvus and A. yahnahpin, to Brontosaurus, as well as the type species B. excelsus. Paleontologist Michael D'Emic made a critique. Paleontologist Donald Prothero criticized the mass media reaction to this study as superficial and premature, concluding:
Until someone has convincingly addressed the issue, I'm going to put "Brontosaurus" in quotes and not follow the latest media fad, nor will I overrule Riggs (1903) and put the name in my books as a valid genus.[1]
Classification[]
Brontosaurus is a member of the family Diplodocidae, a clade of gigantic sauropod dinosaurs. The family includes some of the longest and largest creatures ever to walk the earth, including Diplodocus, Supersaurus, and Barosaurus. Brontosaurus is also classified in the subfamily Apatosaurinae, which also includes Apatosaurus and one or more possible unnamed genera. Othniel Charles Marsh described Brontosaurus as being allied to Atlantosaurus, within the now defunct group Atlantosauridae. In 1878, Marsh raised his family to the rank of suborder, including Apatosaurus, Brontosaurus, Atlantosaurus, Morosaurus (=Camarasaurus), and Diplodocus. He classified this group within Sauropoda. In 1903, Elmer S. Riggs mentioned that the name Sauropoda would be a junior synonym of earlier names, and grouped Apatosaurus within Opisthocoelia. Most authors still use Sauropoda as the group name.
Originally named by its discoverer Othniel Charles Marsh in 1879, Brontosaurus had long been considered a junior synonym of Apatosaurus; its type species, Brontosaurus excelsus, was reclassified as A. excelsus in 1903. However, an extensive study published in 2015 by a joint British-Portuguese research team concluded that Brontosaurus was a valid genus of sauropod distinct from Apatosaurus. Nevertheless, not all paleontologists agree with this division. The same study classified two additional species that had once been considered Apatosaurus and Eobrontosaurus as Brontosaurus parvus and Brontosaurus yahnahpin respectively. Cladogram of the Diplodocidae after Tschopp, Mateus, and Benson (2015):
| Diplodocidae |
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Species[]
- Brontosaurus excelsus, the type species of Brontosaurus, was first named by Marsh in 1879. Many specimens, including the holotype specimen YPM 1980, have been assigned to the species. They include FMNH P25112, the skeleton mounted at the Field Museum of Natural History, which has since been found to represent an unknown species of apatosaurine. Brontosaurus amplus, occasionally assigned to B. parvus, is a junior synonym of B. excelsus. B. excelsus therefore only includes its type specimen and the type specimen of B. amplus. The largest of these specimens is estimated to have weighed up to 15 tonnes and measured up to 22 m (72 ft) long from head to tail. Both known definitive B. excelsus fossils have been reported from Reed's Quarry 10 of the Morrison Formation Brushy Basin member in Albany County, Wyoming, dated to the late Kimmeridgian age, about 152 million years ago.
- Brontosaurus parvus, first described as Elosaurus in 1902 by Peterson and Gilmore, was reassigned to Apatosaurus in 1994, and to Brontosaurus in 2015. Specimens assigned to this species include the holotype, CM 566 (a partial skeleton of a juvenile found in Sheep Creek Quarry 4 in Albany County, WY), BYU 1252-18531 (a nearly complete skeleton found in Utah and mounted at Brigham Young University), and the partial skeleton UW 15556 (which had once been accidentally mixed together with the holotype). It dates to the middle Kimmeridgian. Adult specimens are estimated to have weighed up to 14 tonnes and measured up to 22 m (72 ft) long from head to tail.
- Brontosaurus yahnahpin is the oldest species, known from a single site from the lower Morrison Formation, Bertha Quarry, in Albany County, Wyoming, dating to about 155 million years ago. It grew up to 21 m (69 ft) long. The type species, E. yahnahpin, was described by James Filla and Patrick Redman in 1994 as a species of Apatosaurus (A. yahnahpin). The specific name is derived from Lakota mah-koo yah-nah-pin, "breast necklace", a reference to the pairs of sternal ribs that resemble the hair pipes traditionally worn by the tribe. The holotype specimen is TATE-001, a relatively complete postcranial skeleton found in Wyoming, in the lower Morrison Formation. More fragmentary remains have also been referred to the species. A re-evaluation by Robert T. Bakker in 1998 found it to be more primitive, so Bakker coined the new generic name Eobrontosaurus, derived from Greek eos, "dawn", and Brontosaurus.
The cladogram below is the result of an analysis by Tschopp, Mateus, and Benson (2015). The authors analyzed most diplodocid type specimens separately to deduce which specimen belonged to which species and genus.
| Apatosaurinae |
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Paleobiology[]
Gallery[]
References[]
- ^ Prothero, D. 2015.