|Holotype vertebrae of A. altus, AMNH|
Amphicoelias (//, meaning "biconcave", from the Greek ἀμφί, amphi: "on both sides", and κοῖλος, koilos: "hollow, concave") is a genus of herbivorous sauropod dinosaur, with one known species, A. altus. The scant but exceptionally large remains that were originally believed to represent a second species, A. fragillimus, were reassigned to the new genus Maraapunisaurus in a 2018 study.
The type species of Amphicoelias, A. altus, was named by paleontologist Edward Drinker Cope in December 1877 (though not published until 1878) for an incomplete skeleton consisting of two vertebrae, a pubis (hip bone), and a femur (upper leg bone). Cope also named a second species, A. latus, in the same paper. In 1881, Othniel Charles Marsh concluded that A. altus could not be distinguished from other genera, as the features described by Cope were misinterpreted and are widespread. In 1921, Osborn and Mook assigned additional bones to A. altus—a scapula (shoulder blade), a coracoid (shoulder bone), an ulna (lower arm bone), and a tooth. Henry Fairfield Osborn and Charles Craig Mook noted the overall close similarity between Amphicoelias and Diplodocus, as well as a few key differences, such as proportionally longer forelimbs in Amphicoelias than in Diplodocus. The femur of Amphicoelias is unusually long, slender, and round in cross section; while this roundness was once thought to be another distinguishing characteristic of Amphicoelias, it has since been found in some specimens of Diplodocus as well. A. altus was also similar in size to Diplodocus, estimated to be about 25 m (82 ft) long. Foster (2007) suggested that Amphicoelias is probably the senior synonym of Diplodocus. Whitlock (2011) and Mannion et al. (2012) rejected the referral of the tooth to Amphicoelias as arbitrary, while questioning the referral of the pectoral girdle and ulna to the genus.
Classification and species
Edward Drinker Cope described his finds in two 1878 issues of the American Naturalist, and assigned them to the new genus Amphicoelias. He placed it in a unique family, Amphicoeliidae, though this is now considered a nomen oblitum (forgotten name). The genus is usually assigned to the family Diplodocidae, though some modern analyses have found it at the base of the larger group Diplodocoidea or as a diplodocid incertae sedis (uncertain placement). The first named species in the genus, Amphicoelias altus (holotype specimen AMHD 5764), was discovered by Aaron Ripley, the brother-in-law of Cope's collector Oramel William Lucas, in 1877. But while it is only represented by a partial skeleton, there are enough diagnostic characteristics to provisionally define the genus. A. altus is known from better remains, but is smaller than A. fragillimus. Cope also named a second species in 1878: Amphicoelias latus.
The third named Amphicoelias species, A. fragillimus, was known only from a single, incomplete 1.5 m tall neural arch (the part of a vertebra with spines and processes), either last or second to last in the series of back vertebrae, D (dorsal) 10 or D9. Based only on an illustration published in 1878, this vertebra would have measured 2.7 meters (8.9 ft) tall in life. However, it has been argued that the scale bar in the published description contained a typographical error, and the fossil vertebra was in fact only 1.38 meters (4.5 ft) tall. In addition to this vertebra, Cope's field notes contain an entry for an "[i]mmense distal end of femur”, located only a few tens of meters away from the giant vertebra. It is likely that this undescribed leg bone belonged to the same individual animal as the neural spine. In 2018, "A." fragillimus was given its own genus, Maraapunisaurus.
In 2010, an article was made available, but not formally published, by Henry Galiano and Raimund Albersdorfer in which they referred to the new species "A. brontodiplodocus" to Amphicoelias, based on several complete specimens found in the Dana Quarry of Big Horn Basin, Wyoming and held in a private collection. The specific name referred to their hypothesis based on these specimens that nearly all Morrison diplodocid species are either growth stages or represent sexual dimorphism among members of the genus Amphicoelias, but this analysis has been met with skepticism and the publication itself has been disclaimed by its lead author, explaining that it is "obviously a drafted manuscript complete with typos, etc., and not a final paper. In fact, no printing or distribution has been attempted".
Osborn and Mook, in 1921, provisionally synonymized A. fragillimus with A. altus, while sinking A. latus into Camarasaurus supremus, and suggesting also that A. fragillimus is just a very large individual of A. altus, a position that most subsequent studies, including McIntosh 1998, Foster (2007), and Woodruff and Foster (2015) have agreed with. Carpenter (2006) disagreed about the synonymy of A. altus and A. fragillimus, however, citing numerous differences in the construction of the vertebra also noted by Cope, and suggested these differences are enough to warrant a separate species or even a separate genus for A. fragillimus. However, he went on to caution that the validity of A. fragillimus as a separate species is nearly impossible to determine without the original specimen to study. Although Amphicoelias latus is clearly not Amphicoelias, it is probably synonymous with Camarasaurus grandis rather than C. supremus because it was found lower in the Morrison Formation and the deeply concave articular faces on the caudal vertebrae are more consistent with C. grandis.
In 2007, John Foster suggested that the differences usually cited to differentiate Amphicoelias altus from the more well known Diplodocus are not significant and may be due to individual variation. Foster argued that Amphicoelias is probably the senior synonym of Diplodocus, and that if further research bears this out, the familiar name Diplodocus would need to be abandoned in favor of Amphicoelias, as was the case with Brontosaurus and its senior synonym Apatosaurus. In 2015, Woodruff and Foster reiterated this conclusion, stating that there is only one species of Amphicoelias and that it could be referred to Diplodocus as Diplodocus altus. They considered the name Amphicoelias to be a nomen oblitum. It has also been hypothesized that Amphicoelias should be considered an apatosaurine, and therefore should be placed in the subfamily Apatosaurinae.
In his 2006 re-evaluation, Carpenter examined the paleobiology of giant sauropods, including Amphicoelias, and addressed the question of why this group attained such a huge size. He pointed out that gigantic sizes were reached early in sauropod evolution, with very large sized species present as early as the late Triassic Period, and concluded that whatever evolutionary pressure caused large size was present from the early origins of the group. Carpenter cited several studies of giant mammalian herbivores, such as elephants and rhinoceros, which showed that larger size in plant-eating animals leads to greater efficiency in digesting food. Since larger animals have longer digestive systems, food is kept in digestion for significantly longer periods of time, allowing large animals to survive on lower-quality food sources. This is especially true of animals with a large number of 'fermentation chambers' along the intestine, which allow microbes to accumulate and ferment plant material, aiding digestion. Throughout their evolutionary history, sauropod dinosaurs were found primarily in semi-arid, seasonally dry environments, with a corresponding seasonal drop in the quality of food during the dry season. The environment of Amphicoelias was essentially a savanna, similar to the arid environments in which modern giant herbivores are found, supporting the idea that poor-quality food in an arid environment promotes the evolution of giant herbivores. Carpenter argued that other benefits of large size, such as relative immunity from predators, lower energy expenditure, and longer life span, are probably secondary advantages.
The Morrison Formation environment in which Amphicoelias lived would have resembled a modern savanna, though since grass did not appear until the Late Cretaceous, ferns were probably the dominant plant and main food source for Amphicoelias. Though Engelmann et al. (2004) dismissed ferns as a sauropod food source due to their relatively low caloric content, Carpenter argued that the sauropod digestive system, well adapted to handle low-quality food, allows for the consumption of ferns as a large part of the sauropod diet. Carpenter also noted that the occasional presence of large petrified logs indicate the presence of 20–30 m (66–98 ft) tall trees, which would seem to conflict with the savanna comparison. However, the trees are rare, and since tall trees require more water than the savanna environment could generally provide, they probably existed in narrow tracts or "gallery forests" along rivers and gulleys where water could accumulate. Carpenter speculated that giant herbivores like Amphicoelias may have used the shade of the gallery forests to stay cool during the day, and done most of their feeding on the open savanna at night.
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- Carpenter, Kenneth (2018). "Maraapunisaurus fragillimus, N.G. (formerly Amphicoelias fragillimus), a basal Rebbachisaurid from the Morrison Formation (Upper Jurassic) of Colorado". Geology of the Intermountain West. 5: 227–244.
- Galiano, H. and Albersdorfer, R. "A new basal diplodocid species, Amphicoelias brontodiplodocus, from the Morrison Formation, Big Horn Basin, Wyoming, with taxonomic reevaluation of Diplodocus, Apatosaurus, and other genera. Archived July 10, 2011, at the Wayback Machine." Dinosauria International, LLC. 44pp. Published online 2010.
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- Engelmann, G.F., Chure, D.J., and Fiorillo, A.R. (2004). "The implications of a dry climate for the paleoecology of the fauna of the Upper Jurassic Morrison Formation." In Turner, C.E., Peterson, F., and Dunagan, S.P., eds., Reconstruction of the extinct ecosystem of the Upper Jurassic Morrison Formation: Sedimentary Geology, 167: 297–308
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