Apatosaurus
Apatosaurus is a genus of herbivorous sauropod dinosaur that lived in North America during the Late Jurassic period. Othniel Charles Marsh described and named the first-known species, Apatosaurus ajax, in 1877, and a second species, Apatosaurus louisae, was discovered and named by William H. Holland in 1916. Apatosaurus lived about 152 to 151 million years ago, during the late Kimmeridgian to early Tithonian age, and are now known from fossils in the Morrison Formation of modern-day Colorado, Oklahoma, New Mexico, Wyoming, and Utah in the United States. Apatosaurus had an average length of, and an average mass of. A few specimens indicate a maximum length of 11–30% greater than average and a mass of approximately.
Apatosaurus is a member of the diplodocid family, differing from Diplodocus proper in primarily robustness in the cervical vertebrae and the leg bones. The cervical vertebrae of Apatosaurus are less elongated and more heavily constructed than those of Diplodocus and the bones of the leg are much stockier despite being longer, implying that Apatosaurus was a more robust animal. The tail was held above the ground during normal locomotion. Apatosaurus had a single claw on each forelimb and three on each hindlimb. The Apatosaurus skull, long thought to be similar to Camarasaurus, is much more similar to that of Diplodocus. Apatosaurus was a generalized browser that likely held its head elevated. To lighten its vertebrae, Apatosaurus had air sacs that made the bones internally full of holes. Like that of other diplodocids, its tail may have been used as a whip to create loud noises, or, as more recently suggested, as a sensory organ.
The skull of Apatosaurus was confused with that of Camarasaurus and Brachiosaurus until 1909, when the holotype of A. louisae was found, and a complete skull just a few meters away from the front of the neck. Henry Fairfield Osborn disagreed with this association, and went on to mount a skeleton of Apatosaurus with a Camarasaurus skull cast. Apatosaurus skeletons were mounted with speculative skull casts until 1970, when McIntosh showed that more robust skulls assigned to Diplodocus were more likely from Apatosaurus.
Apatosaurus is a genus in the family Diplodocidae. It is one of the more basal genera, with only Amphicoelias and possibly a new, unnamed genus more primitive. Although the subfamily Apatosaurinae was named in 1929, the group was not used validly until an extensive 2015 study. Only Brontosaurus is also in the subfamily, with the other genera being considered synonyms or reclassified as diplodocines. Brontosaurus has long been considered a junior synonym of Apatosaurus; its type species was reclassified as A.excelsus in 1903. A 2015 study concluded that Brontosaurus is a valid genus of sauropod distinct from Apatosaurus, but not all paleontologists agree with this division. As it existed in North America during the late Jurassic, Apatosaurus would have lived alongside dinosaurs such as Allosaurus, Camarasaurus, Diplodocus, and Stegosaurus.
Description
Apatosaurus was a large, long-necked, quadrupedal animal with a long, whip-like tail. Its forelimbs were slightly shorter than its hindlimbs. Most size estimates are based on specimen CM3018, the type specimen of A.louisae, reaching in length and in body mass. A 2015 study that estimated the mass of volumetric models of Dreadnoughtus, Apatosaurus, and Giraffatitan estimates CM3018 at, similar in mass to Dreadnoughtus. Some specimens of A.ajax represent individuals 1130% longer, suggesting masses twice that of CM3018 or, potentially rivaling the largest titanosaurs. However, the upper size estimate of OMNH1670 is likely an exaggeration, with the size estimates revised in 2020 at in length and in body mass based on volumetric analysis.The skull is small in relation to the size of the animal. The jaws are lined with spatulate teeth suited to an herbivorous diet. The snout of Apatosaurus and similar diplodocoids is squared, with only Nigersaurus having a squarer skull. The braincase of Apatosaurus is well preserved in specimen BYU17096, which also preserved much of the skeleton. A phylogenetic analysis found that the braincase had a morphology similar to those of other diplodocoids. Some skulls of Apatosaurus have been found still in articulation with their teeth. Those teeth that have the enamel surface exposed do not show any scratches on the surface; instead, they display a sugary texture and little wear.
Like those of other sauropods, the neck vertebrae are deeply bifurcated; they carried neural spines with a large trough in the middle, resulting in a wide, deep neck. The vertebral formula for the holotype of A.louisae is 15cervicals, 10dorsals, 5sacrals, and 82caudals. The caudal vertebra number may vary, even within species. The cervical vertebrae of Apatosaurus and Brontosaurus are stouter and more robust than those of other diplodocids and were found to be most similar to Camarasaurus by Charles Whitney Gilmore. In addition, they support cervical ribs that extend farther towards the ground than in diplodocines, and have vertebrae and ribs that are narrower towards the top of the neck, making the neck nearly triangular in cross-section. In Apatosaurus louisae, the atlas-axis complex of the first cervicals is nearly fused. The dorsal ribs are not fused or tightly attached to their vertebrae and are instead loosely articulated. Apatosaurus has ten dorsal ribs on either side of the body. The large neck was filled with an extensive system of weight-saving air sacs. Apatosaurus, like its close relative Supersaurus, has tall neural spines, which make up more than half the height of the individual bones of its vertebrae. The shape of the tail is unusual for a diplodocid; it is comparatively slender because of the rapidly decreasing height of the vertebral spines with increasing distance from the hips. Apatosaurus also had very long ribs compared to most other diplodocids, giving it an unusually deep chest. As in other diplodocids, the tail transformed into a whip-like structure towards the end.
The limb bones are also very robust. Within Apatosaurinae, the scapula of Apatosaurus louisae is intermediate in morphology between those of A.ajax and Brontosaurus excelsus. The arm bones are stout, so the humerus of Apatosaurus resembles that of Camarasaurus, as well as Brontosaurus. However, the humeri of Brontosaurus and A.ajax are more similar to each other than they are to A.louisae. In 1936, Charles Gilmore noted that previous reconstructions of Apatosaurus forelimbs erroneously proposed that the radius and ulna could cross; in life they would have remained parallel. Apatosaurus had a single large claw on each forelimb, a feature shared by all sauropods more derived than Shunosaurus. The first three toes had claws on each hindlimb. The phalangeal formula is 2-1-1-1-1, meaning the innermost finger on the forelimb has two bones and the next has one. The single manual claw bone is slightly curved and squarely truncated on the anterior end. The pelvic girdle includes the robust ilia, and the fused pubes and ischia. The femora of Apatosaurus are very stout and represent some of the most robust femora of any member of Sauropoda. The tibia and fibula bones are different from the slender bones of Diplodocus but are nearly indistinguishable from those of Camarasaurus. The fibula is longer and slenderer than the tibia. The foot of Apatosaurus has three claws on the innermost digits; the digit formula is 3-4-5-3-2. The first metatarsal is the stoutest, a feature shared among diplodocids.
Discovery and species
Initial discovery
The first Apatosaurus fossils were discovered by Arthur Lakes, a local miner, and his friend Henry C. Beckwith in the spring of 1877 in Morrison, a town in the eastern foothills of the Rocky Mountains in Jefferson County, Colorado. Arthur Lakes wrote to Othniel Charles Marsh, Professor of Paleontology at Yale University, and Edward Drinker Cope, a paleontologist based in Philadelphia, about the discovery until eventually collecting several fossils and sending them to both paleontologists. Marsh named Atlantosaurus montanus based on some of the fossils sent and hired Lakes to collect the rest of the material at Morrison and send it to Yale, while Cope attempted to hire Lakes as well but was rejected. One of the best specimens collected by Lakes in 1877 was a well preserved partial postcranial skeleton, including many vertebrae, and a partial braincase, which was sent to Marsh and named Apatosaurus ajax in November 1877. The composite term Apatosaurus comes from the Greek words apatē /apatēlos meaning "deception"/"deceptive", and sauros meaning "lizard"; thus, "deceptive lizard". Marsh gave it this name based on the chevron bones, which are dissimilar to those of other dinosaurs; instead, the chevron bones of Apatosaurus showed similarities with those of mosasaurs, most likely that of the representative species Mosasaurus. By the end of excavations at Lakes' quarry in Morrison, several partial specimens of Apatosaurus had been collected, but only the type specimen of A. ajax can be confidently referred to the species.During excavation and transportation, the bones of the holotype skeleton were mixed with those of another Apatosaurine individual originally described as Atlantosaurus immanis; as a consequence, some elements cannot be ascribed to either specimen with confidence. Marsh distinguished the new genus Apatosaurus from Atlantosaurus on the basis of the number of sacral vertebrae, with Apatosaurus possessing three and Atlantosaurus four. Recent research shows that traits usually used to distinguish taxa at this time were actually widespread across several taxa, causing many of the taxa named to be invalid, like Atlantosaurus. Two years later, Marsh announced the discovery of a larger and more complete specimen from Como Bluff, Wyoming, he gave this specimen the name Brontosaurus excelsus. Also at Como Bluff, the Hubbell brothers working for Edward Drinker Cope collected a tibia, fibula, scapula, and several caudal vertebrae along with other fragments belonging to Apatosaurus in 1877–78 at Cope's Quarry 5 at the site. Later in 1884, Othniel Marsh named Diplodocus lacustris based on a chimeric partial dentary, snout, and several teeth collected by Lakes in 1877 at Morrison. In 2013, it was suggested that the dentary of D. lacustris and its teeth were actually from Apatosaurus ajax based on its proximity to the type braincase of A. ajax. All specimens currently considered Apatosaurus were from the Morrison Formation, the location of the excavations of Marsh and Cope.