Brachiosaurus: Difference between revisions

Brachiosaurus; Temporal range: Late Jurassic, 150–145 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Bronze cast of B. altithorax outside the Field Museum of Natural History
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Superorder:	Dinosauria
Order:	Saurischia
Suborder:	†Sauropodomorpha
Infraorder:	†Sauropoda
Family:	†Brachiosauridae
Genus:	†Brachiosaurus; Riggs, 1903
Species
	B. altithorax Riggs, 1903 (type); __________________________ invalid species "B." atalaiensis de Lapparent and Zbyszewski, 1960, is now considered a separate genus, Lusotitan; "B." brancai Janensch, 1914 is now considered a separate genus, Giraffatitan; "B." fraasi Janensch, 1914 is a junior synonym of Giraffatitan brancai; "B." nougaredi de Lapparent, 1960 is now considered to represent an unnamed genus of brachiosaurid;

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Brachiosaurus is a genus of sauropod dinosaur from the Jurassic Morrison Formation of North America. It was first described by Elmer Riggs in 1903 from fossils found in the Grand River Canyon (now Colorado River) of western Colorado, in the United States. Riggs named the dinosaur Brachiosaurus altithorax, declaring it "the largest known dinosaur". Brachiosaurus had a proportionally long neck, small skull, and large overall size, all of which are typical for sauropods. Also, the limbs were pillar-like, and the hands columnar. However, the proportions of Brachiosaurus are unlike most sauropods. The forelimbs were longer than the hindlimbs, which result in a steeply inclined trunk, making the overall body shape reminiscent of a modern giraffe. Also, while the tail is a typical long dinosaur tail, it was relatively short for a sauropod.

Much of what is known by laypeople about Brachiosaurus is in fact based on Giraffatitan brancai, a species of brachiosaurid dinosaur from the Tendaguru Formation of Tanzania described by German paleontologist Werner Janensch as a species of Brachiosaurus. Recent research shows that the differences between the type species of Brachiosaurus and the Tendaguru material are significant enough that the African material should be placed in a separate genus.

Description

Like all sauropod dinosaurs, Brachiosaurus was a quadrupedal animal with a small skull, a long neck, a large trunk with a high-ellipsoid cross section, a long, muscular tail and slender, columnar limbs.^[1] The skull had a robust, wide muzzle and thick jaw bones, with spoon–shaped teeth. As in Giraffatitan, there was an arch of bone over the snout and in front of the eyes that encircled the nasal opening, although this arch was not as large as in its relative.^[2] Large air sacs connected to the lung system were present in the neck and trunk, invading the vertebrae and ribs, greatly reducing the overall density.^[3]^[4] Unusually for a sauropod, the forelimbs were longer than the hind limbs. The humerus (upper arm bone) of Brachiosaurus was relatively lightly built for its size,^[5] measuring 2.04 metres (6.7 ft) in length in the type specimen.^[6] The femur (thigh bone) of the type specimen was only 2.03 metres (6.7 ft) long.^[6] Unlike other sauropods, Brachiosaurus appears to have been slightly sprawled at the shoulder joint,^[5] and the ribcage was unusually deep.^[6] This led to the trunk being inclined, with the front much higher than the hips, and the neck exiting the trunk at a steep angle. Overall, this shape resembles a giraffe more than any other living animal.^[7]

Size

Because "Brachiosaurus" brancai (Giraffatitan) is known from much more complete material than B. altithorax, most size estimates for Brachiosaurus are actually for the African form. There is an additional element of uncertainty for North American Brachiosaurus because the most complete skeleton appears to have come from a subadult.^[5] Over the years, the mass of B. altithorax has been estimated as 35.0 metric tons (38.6 short tons),^[7] 43.9 metric tons (48.4 short tons),^[8] and, most recently, 28.7 metric tons (31.6 short tons).^[5] In the first and last cases, the authors also provided estimates for Giraffatitan, and found that genus to be somewhat lighter (31.5 metric tons (34.7 short tons) for Paul [1988]^[7] and 23.3 metric tons (25.7 short tons) for Taylor [2009]^[5]). The length of Brachiosaurus has been estimated at 26 metres (85 ft).^[9]

Classification

Brachiosaurus is the namesake genus of Brachiosauridae.^[10] Over the years a number of sauropods have been assigned to Brachiosauridae, such as Astrodon, Bothriospondylus, Dinodocus, Pelorosaurus, Pleurocoelus, and Ultrasaurus,^[11] but most of these are currently regarded as dubious or of uncertain placement.^[1] A phylogenetic analysis of sauropods published in the description of Abydosaurus found that genus to form a clade with Brachiosaurus and Giraffatitan (included in Brachiosaurus).^[12] A more recent analysis focused on possible Asian brachiosaurid material found a clade including Abydosaurus, Brachiosaurus, Cedarosaurus, Giraffatitan, and Paluxysaurus, but not Qiaowanlong, the putative Asian brachiosaurid.^[13] Related genera include Lusotitan and Sauroposeidon.^[1] Brachiosauridae is situated at the base of Titanosauriformes, a group of sauropods that also includes the titanosaurs.^[13]

According to the revised diagnosis by Taylor Brachiosaurus altithorax is diagnosed by a plethora of characters, many to be found on the dorsal (back) vertebrae.^[5] Among the characters playing it in the family Brachiosauridae are a ratio of humerus to femur length greater or equal to 0.9 (i.e., the upper arm bone is at least nearly as long as the thigh bone), and a very flattened femur shaft (ratio ≥1.85).^[5]

Discovery and history

The genus Brachiosaurus, and type species B. altithorax, are based on a partial postcranial skeleton from Fruita, in the valley of the Colorado River of western Colorado.^[14] This specimen was collected from rocks of the Brushy Basin Member of the Morrison Formation^[15] in 1900 by Elmer S. Riggs and his crew from the Field Columbian Museum (now the Field Museum of Natural History) of Chicago.^[6] It is currently cataloged as FMNH P 25107.^[5] Riggs and company were working in the area as a result of favorable correspondence between Riggs and S. M. Bradbury, a dentist in nearby Grand Junction. In 1899 Riggs had sent inquiries to rural locations in the western United States concerning fossil finds, and Bradbury, an amateur collector himself, reported that dinosaur bones had been collected in the area since 1885.^[14] It was Riggs' field assistant H. W. Menke who found FMNH P 25107,^[6] on July 4, 1900, on a small hill later known as Riggs Hill.^[14] Riggs published a short report in 1901, noting the unusual length of the humerus compared to the femur and the extreme overall size and the resulting giraffe-like proportions, as well as the lesser development of the tail, but did not publish a name for the new dinosaur.^[16] The titles of Riggs (1901) and (1903) suggested that the specimen was the largest known dinosaur.^[16]^[6] Riggs followed his 1903 publication that named Brachiosaurus altithorax^[6] with a more detailed description in a monograph in 1904.^[10]

The Fruita skeleton was not the first discovery of Brachiosaurus bones, although it was the first to be recognized as belonging to a new and distinct animal. In 1883, a sauropod skull was found near Garden Park, Colorado, at Felch Quarry 1, and was sent to Othniel Charles Marsh (of "Bone Wars" fame).^[5] Marsh incorporated the skull into his skeletal restoration of "Brontosaurus" (now Apatosaurus).^[5]^[17] It eventually became part of the collections of the National Museum of Natural History, as USNM 5730.^[5] In the 1970s, when Jack McIntosh and David Berman were working on the issue of the true skull of Apatosaurus, they reevaluated the Garden Park skull as more similar to Camarasaurus.^[18] It was described and recognized as a Brachiosaurus skull in 1998 by Kenneth Carpenter and Virginia Tidwell, intermediate in form between Camarasaurus and Giraffatitan brancai (then still considered to be B. brancai).^[19] Because there are no overlapping parts between this skull and FMNH P 25107, it cannot be confidently assigned to a species,^[19]^[5] so it is classified as Brachiosaurus sp.^[19]

Additional discoveries of Brachiosaurus material in North America have been uncommon and consist of a handful of bones. Material has been described from Colorado,^[20]^[21]^[22]^[5] Oklahoma,^[23]^[5] Utah,^[20]^[5] and Wyoming,^[8] ^[5]and undescribed material has been mentioned from several other sites.^[15]^[5] One of these specimens, a shoulder blade from Dry Mesa Quarry, Colorado, is one of the specimens at the center of the Supersaurus/Ultrasauros issue of the 1980s and 1990s. In 1985, James A. Jensen described disarticulated sauropod remains from the quarry as belonging to several taxa, including the new genera Supersaurus and Ultrasaurus,^[24] the latter renamed Ultrasauros shortly thereafter because another sauropod already had the name.^[25] Later study showed that the "ultrasaur" material mostly belonged to Supersaurus, although the shoulder blade did not. Because the holotype of Ultrasauros, a back vertebra, was one of the specimens that was actually from Supersaurus, the name Ultrasauros is a synonym of Supersaurus. The shoulder blade is now assigned to Brachiosaurus, but the species is uncertain.^[21]^[5] In addition, the Dry Mesa "ultrasaur" was not as large as had been thought; the dimensions of the shoulder's coracoid bone indicate that the animal was smaller than Riggs' original specimen of Brachiosaurus.^[5]

Etymology

Riggs derived the genus name from the Greek brachion/βραχιων meaning "arm" and sauros/σαυρος meaning "lizard", because he realized that the length of the arms was unusual for a sauropod.^[6] The species epithet "altithorax" was chosen because of the unusually deep and wide chest cavity, from Latin altus meaning "deep" and Greek thorax/θώραξ (Latin thorax), meaning "breastplate, cuirass, corslet".^[26]

Material and species

Brachiosaurus altithorax

FMNH P 25107, the holotype of both the genus Brachiosaurus and the species B. altithorax, consists of the right humerus (upper arm bone), the right femur (thigh bone), the right ilium (a hip bone), the right coracoid (a shoulder bone), the sacrum (fused vertebrae of the hip), the last seven thoracic (trunk) and two caudal (tail) vertebrae, and a number of ribs.^[5]^[6]^[16] Riggs described the coracoid as from the left side of the body,^[6]^[16]^[10] but restudy has shown it to be a right coracoid.^[5]

Type specimens of other assigned species

"B." atalaiensis: Originally described by de Lapparent and Zbyszewski,^[27] this material's reference to Brachiosaurus was doubted by Upchurch, Barret and Dodson,^[1] who listed it as an unnamed brachiosaurid, and placed in its own genus Lusotitan by Antunes and Mateus.^[28] De Lapparent and Zbyszewski described a series of remains but did not designate a type specimen. Antunes and Mateus selected a partial postcranial skeleton (MIGM 4978, 4798, 4801–4810, 4938, 4944, 4950, 4952, 4958, 4964–4966, 4981–4982, 4985, 8807, 8793–87934) as a lectotype; this specimen includes 28 vertebrae, chevrons, ribs, a possible shoulder blade, humeri, forearm bones, partial left pelvis, lower leg bones, and part of the right ankle. The low neural spines, the prominent deltopectoral crest of the humerus (a muscle attachment site on the upper arm bone), the elongated humerus (very long and slender), and the long axis of the ilium tilted upward indicate that Lusotitan is a brachiosaurid.^[28]

"B." brancai: Janensch based his description on "Skelett S" (skeleton S) from Tendaguru,^[29] but later realized that it comprised two partial individuals: S I and S II.^[30] He did not designate them as a syntype series, nor specify a lectotype, and Taylor proposed the larger and more complete S II (MB.R.2181) as the lectotype.^[5] It includes, among other bones, several dorsal (trunk) vertebrae, the left scapula, both coracoids , both sternals (breastbones), both humeri, both ulna and radii (lower arm bones), a right hand, a partial left hand, both pubes (a hip bone) and the right femur, tibia and fibula (shank bones). A re-assessment of the relation between the African and American brachiosaur material indicates that a separate generic name is warranted for the Tendaguru material, meaning that it now is considered to belong to Giraffatitan.^[7]^[5]

"B." fraasi: erected by Janensch in 1914, but later synonomized with "B." brancai;^[30] this material now belongs to Giraffatitan.^[5]

"B." nougaredi: This species is known from fragmentary remains discovered in eastern Algeria, in the Sahara Desert. The present type material consists of a sacrum and some of the left metacarpals and phalanges. Found at the discovery site but not collected were partial bones of the left forearm, wrist bones, a right shin bone, and fragments that may have come from metatarsals.^[31] de Lapparent, who described and named the material in 1960, reported the discovery locality as being in the Late Jurassic–age Taouratine Series (he assigned the rocks this age in part because of the presumed presence of Brachiosaurus),^[31] but more recent review assigns it to the "Continental intercalaire," which is considered to be of Albian age (late Early Cretaceous, significantly younger).^[1] This material was found disjointed over an area of several hundred meters,^[31] and may not belong to one individual or even one species.^{[citation needed]} Upchurch, Barrett and Dodson (2004) doubted its assignment to Brachiosaurus, and listed it as an unnamed brachiosaurid.^[1] The sacrum is of notable size, with a length of 130 centimetres (51 in) for four vertebrae compared to 95 centimetres (37 in) for five vertebrae in B. altithorax; other bones are not of unusual size in comparison to B. altithorax or Giraffatitan brancai.^[31]

Referred material

Taylor (2009) lists a number of specimens referred to Brachiosaurus. These include some material, e.g., a humerus from Potter Creek and some Dry Mesa material (the latter partly described as Ultrasauros by Jensen), that are either clearly not brachiosaurid in origin, or at least not clearly referable to Brachiosaurus.^[5] In contrast, a cervical (neck) vertebra and the skull mentioned above may belong to either B. altithorax or an as-yet unknown brachiosaurid from North America.^[5] The cervical was found near Jensen, Utah, by Jensen,^[20] and – if it belongs to Brachiosaurus – is one of a handful of neck vertebrae known for American brachiosaurids.^[5] There is no unambiguous material of the skull, neck, anterior dorsal (forward trunk) region, distal (lower) limbs or feet.^[5]

There was ample material referred to "B." brancai in the collections of the Museum für Naturkunde Berlin, some of which was destroyed during World War II. Other material was transferred to other institution throughout Germany, some of which was also destroyed. Additional specimens are likely among the material collected by the British Museum of Natural History's Tendaguru expedition.^[32] Much or all of this material probably belongs to Giraffatitan, although some may represent a new brachiosaurid.^[33]

Separation from Giraffatitan

When describing the brachiosaurid material from Tendaguru in 1914, Janensch listed a number of differences and commonalities between them and B. altithorax.^[29] In three further publications in 1929,^[30] 1950 ^[34] and 1961^[35] Janensch compared the two species in more detail, listing 13 putative shared characters.^[5] Of these, however, only four appear to be valid, while six pertain to more inclusive groups than Brachiosauridae, and the rest are either difficult to assess or refer to material that is not Brachiosaurus.^[5]

In 1988, Gregory Paul published a new reconstruction of the skeleton of "B." brancai, highlighting a number of differences in proportion between it and B. altithorax. Chief among them is a difference in the way the trunk vertebrae vary: they are fairly uniform in B. altithorax, but vary widely in the African material. Paul believed that the limb and girdle elements of both species were very similar, and therefore suggested to separate them not at genus, but only at subgenus level.^[7]

Giraffatitan was raised to genus level by Olshevsky without comment.^[25] A detailed study of all material, including the limb and girdle bones, by Michael Taylor in 2009 found that there are significant differences between Brachiosaurus altithorax and the Tendaguru material in all elements known from both species. Taylor found 26 distinct osteological (bone-based) characters, a larger difference than that between, e.g., Diplodocus and Barosaurus, and therefore argued that the African material should be placed in its own genus, Giraffatitan, as G. brancai.^[5] An important difference between the two genera is the overall body shape, with Brachiosaurus having a 23% longer dorsal (trunk) vertebrate series and a 20 to 25% longer and also taller tail.^[5]

Paleoecology

With the removal of the East African Giraffatitan, Brachiosaurus is known only from the Morrison Formation of western North America.^[5] The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons,^[36]^[37] and flat floodplains.^[36] Vegetation varied from gallery forests (river–lining forests in otherwise treeless settings) of conifers, tree ferns, and ferns, to fern savannas with rare trees.^[38] Several other sauropod genera were present in the Morrison Formation, with differing body proportions and feeding adaptations.^[8] Among these were Apatosaurus, Barosaurus, Camarasaurus, Diplodocus, Haplocanthosaurus, and Supersaurus.^[8]^[39] Brachiosaurus was one of the less abundant Morrison Formation sauropods. In a survey of over 200 fossil localities, John Foster reported 12 specimens of the genus, comparable to Barosaurus (13) and Haplocanthosaurus (12), but far fewer than Apatosaurus (112), Camarasaurus (179), and Diplodocus (98).^[8]

Paleobiology

Neck position

In contrast to most other sauropods, brachiosaurids had an inclined back, due to their long forelimbs. Therefore, if the neck exited the body in a straight line, it already pointed upwards.^[5]^[40]^[41]^[42] The exact angle is influenced by how the pectoral girdle is reconstructed, that is how the shoulder blades are placed on the ribcage.^[40]^[41]^[42] The mobility of the neck was reconstructed as quite low by Stevens and Parrish,^[40]^[41]^[42] while other researchers like Paul and Christian and Dzemski argued for more flexible necks.^[7]^[43]

Feeding ecology

Brachiosaurus is thought to have been a high browser, feeding on foliage well above the ground. Even if it did not hold its neck near vertical, and instead had a straighter neck, its head height may still have been over 9 metres (30 ft) above the ground.^[8]^[2] It probably fed mostly on foliage above 5 metres (16 ft). This does not preclude the possibility that it also fed lower at times, between 3 to 5 metres (9.8 to 16.4 ft) up.^[2] Its diet likely consisted of ginkgoes, conifers, tree ferns, and large cycads, with intake estimated at 200 to 400 kilograms (440 to 880 lb) of plant matter daily.^[2] However, more recent studies estimate that ~240 kilograms (530 lb) of plant matter would have been sufficient to feed a 70 metric tons (77 short tons) sauropod,^[44] so Brachiosaurus may have required only about 120 kilograms (260 lb) of fodder a day. Brachiosaur feeding involved simple up–and–down jaw motion. The teeth were arranged to shear material as they closed, and were probably used to crop and/or nip vegetation.^[45]

It has repeatedly been suggested, e.g. in the movie Jurassic Park, that Brachiosaurus could rear into a bipedal or tripodal (with tail support) pose to feed. However, a detailed physical modelling-based analysis of sauropod rearing capabilities by Heinrich Mallison showed that while many sauropods could rear, the unusual brachiosaurid body shape and limb length ratio made them exceptionally ill suited for rearing. The forward position of the center of mass would have led to problems with stability, and required unreasonably large forces in the hips to obtain an upright posture. Brachiosaurus would also have gained relatively little from rearing (only 33% more feeding height), compared to other sauropods, for which a bipedal pose may have tripled the feeding height.^[46]

Metabolism

Like all sauropods, Brachiosaurus was homeothermic (maintaining a stable internal temperature) and endothermic (controlling body temperature through internal means), meaning that it was able to actively control its body temperature, producing the necessary heat through a high basic metabolic rate of its cells.^[47]

In the past, Brachiosaurus has been used an example of a dinosaur for which endothermy is unlikely, because of the combination of great size (leading to overheating) and great caloric needs to fuel endothermy.^[48] However, these calculations were based on incorrect assumptions about the available cooling surfaces (the large air sacs were not known), and a grossly inflated body mass. These inaccuracies resulted in overestimation of heat production and underestimation of heat loss.^[47] The large nasal arch has been postulated as an adaptation for cooling the brain, as a surface for evaporative cooling of the blood.^[48] Again, when this was proposed, cooling via the air sacs was not known, and thus not taken into account. Furthermore, other similar sized sauropods had no comparable structure. Additionally, in proportion to the entire animal, the nasal arch is very small, and would thus have made only an insignificant contribution to heat loss.

In culture

The original Brachiosaurus specimens collected by Elmer Riggs for the Field Museum of Natural History (Chicago) were not put on display in the museum until 1994, when a skeletal mount (made up of resin casts rather than actual fossil bones) was constructed inside the museum's main Stanley Field Hall. The mount stood until 1999, when it was moved to the B Concourse of United Airlines' Terminal One in O'Hare International Airport.^[49] At the same time, a second cast (in bronze) of the Field Museum's B. altithorax was constructed outside the museum.^[50]

Brachiosaurus is one of the best-known dinosaurs amongst both paleontologists and the general public. A main belt asteroid, 1991 GX₇, has been named 9954 Brachiosaurus in honor of the genus.^[51]^[52] The genus has been featured in many films and television programs, most notably the Jurassic Park and Walking with Dinosaurs series. The digital model of Brachiosaurus used in Jurassic Park went on to become the starting point for the ronto models in the 1997 special edition of the science fiction film Star Wars Episode IV: A New Hope.^[53]

References

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^ ^a ^b ^c ^d Foster, J. (2007). "Brachiosaurus altithorax." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 205–208.
^ Wedel, M.J. (2003). "Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs." Paleobiology 29:243-255.
^ Wedel, M.J. (2003). "The evolution of vertebral pneumaticity in sauropod dinosaurs." Journal of Vertebrate Paleontology 23:344-357.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah Taylor, M.P. (2009). "A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensh 1914)." Journal of Vertebrate Paleontology, 29(3): 787-806.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Riggs, E.S. (1903). "Brachiosaurus altithorax, the largest known dinosaur." American Journal of Science, 4(15): 299-306.
^ ^a ^b ^c ^d ^e ^f Paul, G.S. (1988). "The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world's largest dinosaurs". Hunteria. 2 (3).
^ ^a ^b ^c ^d ^e ^f Foster, J.R. (2003). Paleoecological analysis of the vertebrate fauna of the Morrison Formation (Upper Jurassic), Rocky Mountain region, U.S.A. New Mexico Museum of Natural History and Science Bulletin, 23. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science.
^ Holtz, T.R. Jr. (2008) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages Supplementary Information
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^ Lambert, David (1990). "Brachiosaurids". The Dinosaur Data Book. New York: Avon Books. p. 142. ISBN 0-380-75896-3. {{cite book}}: Check |isbn= value: checksum (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Chure, D. (2010). "First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition" (PDF). Naturwissenschaften. 97 (4): 379–391. doi:10.1007/s00114-010-0650-6. PMC 2841758. PMID 20179896. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b Ksepka, D. T. (2010). "The illusory evidence for Asian Brachiosauridae: new material of Erketu ellisoni and a phylogenetic appraisal of basal Titanosauriformes" (pdf). American Museum Novitates. 3700: 1–27. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c Glut, D.F. (1997). "Brachiosaurus". Dinosaurs: The Encyclopedia. McFarland & Company. pp. 213–221. ISBN 0-89950-917-7.
^ ^a ^b Turner, C.E. (1999). "Biostratigraphy of dinosaurs in the Upper Jurassic Morrison Formation of the Western Interior, USA". In Gillete, David D. (ed.) (ed.). Vertebrate Paleontology in Utah. Miscellaneous Publication 99-1. Salt Lake City, Utah: Utah Geological Survey. pp. 77–114. ISBN 1-55791-634-9. {{cite book}}: |editor= has generic name (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c ^d Riggs, E.S. (1091). "The largest known dinosaur". Science. 13 (327): 549–550. doi:10.1126/science.13.327.549-a. PMID 17801098.
^ Marsh, O.C. (1891). "Restoration of Triceratops" (pdf). American Journal of Science. 41 (244): 339–342.
^ McIntosh, J.S. (1975). "Description of the palate and lower jaw of the sauropod dinosaur Diplodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus". Journal of Paleontology. 49 (1): 187–199. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c Carpenter, K. and Tidwell, V. (1998). "Preliminary description of a Brachiosaurus skull from Felch Quarry 1, Garden Park, Colorado." Pp. 69–84 in: Carpenter, K., Chure, D. and Kirkland, J. (eds.), The Upper Jurassic Morrison Formation: An Interdisciplinary Study. Modern Geology, 23:1-4.
^ ^a ^b ^c Jensen, J.A. (1987). "New brachiosaur material from the Late Jurassic of Utah and Colorado". The Great Basin Naturalist. 47 (4): 592–608.
^ ^a ^b Curtice, B., Stadtman, K., and Curtice, L. (1996) "A re-assessment of Ultrasauros macintoshi (Jensen, 1985)." Pp. 87-95 in M. Morales (ed.), The Continental Jurassic: Transactions of the Continental Jurassic Symposium, Museum of Northern Arizona Bulletin number 60.
^ Curtice, B. (2001). "The demise of Dystylosaurus edwini and a revision of Supersaurus vivianae". In McCord, R.D.; and Boaz, D. (eds.) (ed.). Western Association of Vertebrate Paleontologists and Southwest Paleontological Symposium - Proceedings 2001. Mesa Southwest Museum Bulletin. Vol. 8. pp. 33–40. {{cite book}}: |editor= has generic name (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: multiple names: editors list (link)
^ Bonnan, M.F. (2004). "First occurrence of Brachiosaurus (Dinosauria, Sauropoda) from the Upper Jurassic Morrison Formation of Oklahoma". PaleoBios. 24 (2): 12–21. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Jensen, J.A. (1985). "Three new sauropod dinosaurs from the Upper Jurassic of Colorado". The Great Basin Naturalist. 45 (4): 697–709.
^ ^a ^b Olshevsky, G. (1991). "A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia." Mesozoic Meanderings 2:1-196
^ θώραξ, Henry George Liddell, Robert Scott, A Greek-English Lexicon, on Perseus Digital Library
^ de Lapparent,A.F. & Zbyszewski, G. (1957). "Les dinosauriens du Portugal". Mémoire Service géologique Portugal 2:1–63.
^ ^a ^b Antunes, M. (2003). "Dinosaurs of Portugal". Comptes rendus. Palévol. 2 (1): 77–95. doi:10.1016/S1631-0683(03)00003-4. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b Janensch, W. (1914). "Übersicht über der Wirbeltierfauna der Tendaguru-Schichten nebst einer kurzen Charakterisierung der neu aufgefuhrten Arten von Sauropoden." Archiv fur Biontologie, 3: 81–110.
^ ^a ^b ^c Janensch, W. (1929). "Material und Formengehalt der Sauropoden in der Ausbeute der Tendaguru-Expedition." Palaeontographica (Suppl. 7) 2:1–34.
^ ^a ^b ^c ^d de Lapparent, A.F. (1960): "Les dinosauriens du "continental intercalaire" du Sahara central" ("The dinosaurs of the "continental intercalaire" of the central Sahara.") Mémoires de la Société Géologic de France, Nouvelle Série 88A vol.39(1-6):1-57. [in French; a translated version, by Matthew Carrano (pdf, no figures), is available through the Polyglot Paleontologist]
^ Maier, G. (2003). African dinosaurs unearthed. The Tendaguru Expeditions." Bloomington, IN: Indiana University Press.
^ Taylor, M. (18 Nov. 2009): CT-scanning the Archbishop. Sauropod Vertebrate Picture of the Week (Blog) post at http://svpow.wordpress.com/2009/11/18/ct-scanning-the-archbishop/.
^ Janensch, W. (1950). "Die Wirbelsäule von Brachiosaurus brancai." Palaeontographica (Suppl. 7) 3:27–93.
^ Janensch, W. (1961). "Die Gliedmaßen und Gliedmaßengürtel der Sauropoden der Tendaguru-Schichten." Palaeontographica (Suppl. 7) 3:177–235.
^ ^a ^b Russell, D. A. (1989). An Odyssey in Time: Dinosaurs of North America. Minocqua, Wisconsin: NorthWord Press. pp. 64–70. ISBN 1-55971-038-1.
^ 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." Sedimentary Geology 167(3-4):297-308.
^ Carpenter, K. (2006). "Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus". Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin, 36. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science. pp. 131–138. {{cite book}}: Unknown parameter |editors= ignored (|editor= suggested) (help)
^ Chure, D.J. (2006). "The fauna and flora of the Morrison Formation: 2006". In Foster, J.R.; and Lucas, S.G. (eds.) (ed.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin, 36. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science. pp. 233–248. {{cite book}}: |editor= has generic name (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: multiple names: editors list (link)
^ ^a ^b ^c Stevens, K. A. and Parrish, M. J. (1999). "Neck posture and feeding habits of two Jurassic sauropod dinosaurs." Science 284:798–800. doi:10.1126/science.284.5415.798
^ ^a ^b ^c Stevens, K. A. and Parrish, M. J. (2005). "Digital reconstructions of sauropod dinosaurs and implications for feeding." In The sauropods: evolution and paleobiology (eds. J. A.Wilson & K. Curry-Rogers), pp. 178–200. Berkeley, CA: University of California Press.
^ ^a ^b ^c Stevens, K. A. and Parrish, M. J. (2005). "Neck posture, dentition and feeding strategies in Jurassic sauropod dinosaurs." In Thunder Lizards: The Sauropodomorph dinosaurs" (eds. V. Tidwell & K. Carpenter). Bloomington, IN: Indiana University Press.
^ Dzemski, G. and Christian, A. (2007). "Flexibility along the neck of the ostrich (Struthio camelus) and consequences for the reconstruction of dinosaurs with extreme neck length." Journal of Morphology 268:701-714.
^ Hummel, J., Gee, C.T., Südekum, K.-H., Sander, P.M., Nogge, G. and Clauss, M. (2008). "In vitro digestibility of fern and gymnosperm foliage: implications for sauropod feeding ecology and diet selection". Proceedings of the Royal Society B, 275:1015-1021. doi:10.1098/rspb.2007.1728
^ Barrett, Paul M. (2005). "Sauropodomorph diversity through time". The Sauropods: Evolution and Paleobiology. Berkeley, CA: University of California. pp. 125–156. ISBN 0-520-24623-3. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |editors= ignored (|editor= suggested) (help)
^ Mallison, H. (in press). "Rearing Giants – kinetic-dynamic modeling of sauropod bipedal and tripodal poses." In Klein, N., Remes, K., Gee, C. & Sander M. (eds): Biology of the Sauropod Dinosaurs: Understanding the life of giants. Life of the Past (series ed. Farlow, J.). Bloomington, IN: Indiana University Press.
^ ^a ^b Sander, P.M., Christian, A., Clauss, M., Fechner, R., Gee, C.T., Griebeler, E.-M., Gunga, H.-C., Hummel, J., Mallison, H., Perry, S.F., Preuschoft, H., Rauhut, O.W.M., Remes, K., Tütken, T., Wings, O. & Witzel, U. (2010). Biology of the sauropod dinosaurs: the evolution of gigantism. Biology Reviews online first publication, doi:10.1111/j.1469-185X.2010.00137.x http://www3.interscience.wiley.com/cgi-bin/fulltext/123397084/HTMLSTART
^ ^a ^b Russell, D. A. (1989). An Odyssey in Time: Dinosaurs of North America. Minocqua, Wisconsin: NorthWord Press. p. 78. ISBN 1-55971-038-1.
^ "Expect Awe-Struck Travelers" (Press release). The Field Museum. November 26, 1999. Retrieved 2009-08-27.
^ The Field Museum. "Captions from Selected Historical Photographs (caption number GN89396_52c)." The Field Museum Photo Archives. pdf link. Accessed 2009-Aug-27.
^ "JPL Small-Body Database Browser: 9954 Brachiosaurus (1991 GX7)". NASA. Retrieved 2007-04-28.
^ Williams, G. "Minor Planet Names: Alphabetical List". Smithsonian Astrophysical Observatory. Retrieved 2007-02-10.
^ "Ronto". Databank. Star Wars.com. Retrieved 2009-01-13.

External links

Expect awe-struck travelers, from the Field Museum. (O'Hare airport mount)
Dinosaurier-info, fact-sheet with picture (in German)
Benes, Josef. Prehistoric Animals and Plants: old-fashioned illustration of Brachiosaurus, bottom left.
Why was Brachiosaurus that big? Short summary of a peer review paper
Brachiosaurus at DinoData
dB Brachiosaurus (Natural History Museum)

[upchurch&al2004-1] ^ ^a ^b ^c ^d ^e ^f Upchurch, P., Barrett, P.M. & Dodson, P. (2004): "Sauropoda." Pp. 259-322 in Weishampel, D.B., Dodson, P. and Osmolska, H. (eds.): The Dinosauria, Second Edition. University of California Press, Berkeley. ISBN 9780520242098

[foster2007b-2] Foster, J. (2007). "Brachiosaurus altithorax." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 205–208.

[wedel2003a-3] Wedel, M.J. (2003). "Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs." Paleobiology 29:243-255.

[wedel2003b-4] Wedel, M.J. (2003). "The evolution of vertebral pneumaticity in sauropod dinosaurs." Journal of Vertebrate Paleontology 23:344-357.

[taylor2009-5] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah Taylor, M.P. (2009). "A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensh 1914)." Journal of Vertebrate Paleontology, 29(3): 787-806.

[riggs1903-6] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Riggs, E.S. (1903). "Brachiosaurus altithorax, the largest known dinosaur." American Journal of Science, 4(15): 299-306.

[paul1988-7] ^ ^a ^b ^c ^d ^e ^f Paul, G.S. (1988). "The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world's largest dinosaurs". Hunteria. 2 (3).

[Foster2003-8] ^ ^a ^b ^c ^d ^e ^f Foster, J.R. (2003). Paleoecological analysis of the vertebrate fauna of the Morrison Formation (Upper Jurassic), Rocky Mountain region, U.S.A. New Mexico Museum of Natural History and Science Bulletin, 23. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science.

[holtz2008-9] Holtz, T.R. Jr. (2008) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages Supplementary Information

[riggs1904-10] Riggs, E.S. (1904). "Structure and relationships of opisthocoelian dinosaurs. Part II. The Brachiosauridae". Geological Series (Field Columbian Museum). 2 (6): 229–247.

[lambert1990-11] Lambert, David (1990). "Brachiosaurids". The Dinosaur Data Book. New York: Avon Books. p. 142. ISBN 0-380-75896-3. {{cite book}}: Check |isbn= value: checksum (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[chure&al2010-12] Chure, D. (2010). "First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition" (PDF). Naturwissenschaften. 97 (4): 379–391. doi:10.1007/s00114-010-0650-6. PMC 2841758. PMID 20179896. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[ksepkanorell2010-13] Ksepka, D. T. (2010). "The illusory evidence for Asian Brachiosauridae: new material of Erketu ellisoni and a phylogenetic appraisal of basal Titanosauriformes" (pdf). American Museum Novitates. 3700: 1–27. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[glut1997-14] Glut, D.F. (1997). "Brachiosaurus". Dinosaurs: The Encyclopedia. McFarland & Company. pp. 213–221. ISBN 0-89950-917-7.

[turner&peterson1999-15] Turner, C.E. (1999). "Biostratigraphy of dinosaurs in the Upper Jurassic Morrison Formation of the Western Interior, USA". In Gillete, David D. (ed.) (ed.). Vertebrate Paleontology in Utah. Miscellaneous Publication 99-1. Salt Lake City, Utah: Utah Geological Survey. pp. 77–114. ISBN 1-55791-634-9. {{cite book}}: |editor= has generic name (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[riggs1901-16] Riggs, E.S. (1091). "The largest known dinosaur". Science. 13 (327): 549–550. doi:10.1126/science.13.327.549-a. PMID 17801098.

[marsh1891-17] Marsh, O.C. (1891). "Restoration of Triceratops" (pdf). American Journal of Science. 41 (244): 339–342.

[mcintosh&berman1975-18] McIntosh, J.S. (1975). "Description of the palate and lower jaw of the sauropod dinosaur Diplodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus". Journal of Paleontology. 49 (1): 187–199. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[carp&tid1998-19] Carpenter, K. and Tidwell, V. (1998). "Preliminary description of a Brachiosaurus skull from Felch Quarry 1, Garden Park, Colorado." Pp. 69–84 in: Carpenter, K., Chure, D. and Kirkland, J. (eds.), The Upper Jurassic Morrison Formation: An Interdisciplinary Study. Modern Geology, 23:1-4.

[Jjenson1987-20] Jensen, J.A. (1987). "New brachiosaur material from the Late Jurassic of Utah and Colorado". The Great Basin Naturalist. 47 (4): 592–608.

[curtice&al1996-21] Curtice, B., Stadtman, K., and Curtice, L. (1996) "A re-assessment of Ultrasauros macintoshi (Jensen, 1985)." Pp. 87-95 in M. Morales (ed.), The Continental Jurassic: Transactions of the Continental Jurassic Symposium, Museum of Northern Arizona Bulletin number 60.

[curt&stad2001-22] Curtice, B. (2001). "The demise of Dystylosaurus edwini and a revision of Supersaurus vivianae". In McCord, R.D.; and Boaz, D. (eds.) (ed.). Western Association of Vertebrate Paleontologists and Southwest Paleontological Symposium - Proceedings 2001. Mesa Southwest Museum Bulletin. Vol. 8. pp. 33–40. {{cite book}}: |editor= has generic name (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: multiple names: editors list (link)

[Bonnan&wedel2004-23] Bonnan, M.F. (2004). "First occurrence of Brachiosaurus (Dinosauria, Sauropoda) from the Upper Jurassic Morrison Formation of Oklahoma". PaleoBios. 24 (2): 12–21. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[jenson1985-24] Jensen, J.A. (1985). "Three new sauropod dinosaurs from the Upper Jurassic of Colorado". The Great Basin Naturalist. 45 (4): 697–709.

[olshevsky1991-25] Olshevsky, G. (1991). "A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia." Mesozoic Meanderings 2:1-196

[thoraxdict-26] θώραξ, Henry George Liddell, Robert Scott, A Greek-English Lexicon, on Perseus Digital Library

[lapp&zby1957-27] Lapparent,A.F. & Zbyszewski, G. (1957). "Les dinosauriens du Portugal". Mémoire Service géologique Portugal 2:1–63.

[antunes&mateus2003-28] Antunes, M. (2003). "Dinosaurs of Portugal". Comptes rendus. Palévol. 2 (1): 77–95. doi:10.1016/S1631-0683(03)00003-4. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[janensch1914-29] Janensch, W. (1914). "Übersicht über der Wirbeltierfauna der Tendaguru-Schichten nebst einer kurzen Charakterisierung der neu aufgefuhrten Arten von Sauropoden." Archiv fur Biontologie, 3: 81–110.

[janensch1929-30] Janensch, W. (1929). "Material und Formengehalt der Sauropoden in der Ausbeute der Tendaguru-Expedition." Palaeontographica (Suppl. 7) 2:1–34.

[lapparent1960-31] Lapparent, A.F. (1960): "Les dinosauriens du "continental intercalaire" du Sahara central" ("The dinosaurs of the "continental intercalaire" of the central Sahara.") Mémoires de la Société Géologic de France, Nouvelle Série 88A vol.39(1-6):1-57. [in French; a translated version, by Matthew Carrano (pdf, no figures), is available through the Polyglot Paleontologist]

[maier2003-32] Maier, G. (2003). African dinosaurs unearthed. The Tendaguru Expeditions." Bloomington, IN: Indiana University Press.

[SVPOW2009-11-18-33] Taylor, M. (18 Nov. 2009): CT-scanning the Archbishop. Sauropod Vertebrate Picture of the Week (Blog) post at http://svpow.wordpress.com/2009/11/18/ct-scanning-the-archbishop/.

[janensch1950a-34] Janensch, W. (1950). "Die Wirbelsäule von Brachiosaurus brancai." Palaeontographica (Suppl. 7) 3:27–93.

[janensch1961-35] Janensch, W. (1961). "Die Gliedmaßen und Gliedmaßengürtel der Sauropoden der Tendaguru-Schichten." Palaeontographica (Suppl. 7) 3:177–235.

[russell1989-36] Russell, D. A. (1989). An Odyssey in Time: Dinosaurs of North America. Minocqua, Wisconsin: NorthWord Press. pp. 64–70. ISBN 1-55971-038-1.

[engelmann&al04-37] 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." Sedimentary Geology 167(3-4):297-308.

[carpenter2006-38] Carpenter, K. (2006). "Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus". Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin, 36. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science. pp. 131–138. {{cite book}}: Unknown parameter |editors= ignored (|editor= suggested) (help)

[Dchure&al2006-39] Chure, D.J. (2006). "The fauna and flora of the Morrison Formation: 2006". In Foster, J.R.; and Lucas, S.G. (eds.) (ed.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin, 36. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science. pp. 233–248. {{cite book}}: |editor= has generic name (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: multiple names: editors list (link)

[stevensparrish1999-40] Stevens, K. A. and Parrish, M. J. (1999). "Neck posture and feeding habits of two Jurassic sauropod dinosaurs." Science 284:798–800. doi:10.1126/science.284.5415.798

[stevensparrish2005-41] Stevens, K. A. and Parrish, M. J. (2005). "Digital reconstructions of sauropod dinosaurs and implications for feeding." In The sauropods: evolution and paleobiology (eds. J. A.Wilson & K. Curry-Rogers), pp. 178–200. Berkeley, CA: University of California Press.

[stevensparrish2005b-42] Stevens, K. A. and Parrish, M. J. (2005). "Neck posture, dentition and feeding strategies in Jurassic sauropod dinosaurs." In Thunder Lizards: The Sauropodomorph dinosaurs" (eds. V. Tidwell & K. Carpenter). Bloomington, IN: Indiana University Press.

[dzemskichristian2007-43] Dzemski, G. and Christian, A. (2007). "Flexibility along the neck of the ostrich (Struthio camelus) and consequences for the reconstruction of dinosaurs with extreme neck length." Journal of Morphology 268:701-714.

[hummel&al2008-44] Hummel, J., Gee, C.T., Südekum, K.-H., Sander, P.M., Nogge, G. and Clauss, M. (2008). "In vitro digestibility of fern and gymnosperm foliage: implications for sauropod feeding ecology and diet selection". Proceedings of the Royal Society B, 275:1015-1021. doi:10.1098/rspb.2007.1728

[bar&up05-45] Barrett, Paul M. (2005). "Sauropodomorph diversity through time". The Sauropods: Evolution and Paleobiology. Berkeley, CA: University of California. pp. 125–156. ISBN 0-520-24623-3. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |editors= ignored (|editor= suggested) (help)

[mallisonsaubook2-46] Mallison, H. (in press). "Rearing Giants – kinetic-dynamic modeling of sauropod bipedal and tripodal poses." In Klein, N., Remes, K., Gee, C. & Sander M. (eds): Biology of the Sauropod Dinosaurs: Understanding the life of giants. Life of the Past (series ed. Farlow, J.). Bloomington, IN: Indiana University Press.

[sander&al2010-47] Sander, P.M., Christian, A., Clauss, M., Fechner, R., Gee, C.T., Griebeler, E.-M., Gunga, H.-C., Hummel, J., Mallison, H., Perry, S.F., Preuschoft, H., Rauhut, O.W.M., Remes, K., Tütken, T., Wings, O. & Witzel, U. (2010). Biology of the sauropod dinosaurs: the evolution of gigantism. Biology Reviews online first publication, doi:10.1111/j.1469-185X.2010.00137.x http://www3.interscience.wiley.com/cgi-bin/fulltext/123397084/HTMLSTART

[russell1989b-48] Russell, D. A. (1989). An Odyssey in Time: Dinosaurs of North America. Minocqua, Wisconsin: NorthWord Press. p. 78. ISBN 1-55971-038-1.

[fmnhpr-49] "Expect Awe-Struck Travelers" (Press release). The Field Museum. November 26, 1999. Retrieved 2009-08-27.

[fieldcaptions-50] The Field Museum. "Captions from Selected Historical Photographs (caption number GN89396_52c)." The Field Museum Photo Archives. pdf link. Accessed 2009-Aug-27.

[51] "JPL Small-Body Database Browser: 9954 Brachiosaurus (1991 GX7)". NASA. Retrieved 2007-04-28.

[Williams-52] Williams, G. "Minor Planet Names: Alphabetical List". Smithsonian Astrophysical Observatory. Retrieved 2007-02-10.

[ronto-53] "Ronto". Databank. Star Wars.com. Retrieved 2009-01-13.

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

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@@ Line 32: / Line 32: @@
 }}
-''Brachiosaurus'' is a [[genus]] of [[sauropod]] [[dinosaur]] from the [[Jurassic]] [[Morrison Formation]] of [[North America]]. It was first described by [[Elmer Riggs]] in 1903 from fossils found in the Grand River Canyon (now [[Colorado River]]) of western [[Colorado]], in the [[United States]]. Riggs named the dinosaur ''Brachiosaurus altithorax'', declaring it "the largest known dinosaur". ''Brachiosaurus''  had a proportionally long neck, small skull, and large overall size, all of which are typical for sauropods. Also, the limbs were pillar-like, and the hands columnar. However, the proportions of ''Brachiosaurus'' are unlike most sauropods. The forelimbs were longer than the hindlimbs, which result in a steeply inclined trunk, making the overall body shape reminiscent of a modern [[giraffe]]. Also, while the tail is a typical long dinosaur tail, it was relatively short for a sauropod.
+'''''Brachiosaurus''''' is a [[genus]] of [[sauropod]] [[dinosaur]] from the [[Jurassic]] [[Morrison Formation]] of [[North America]]. It was first described by [[Elmer Riggs]] in 1903 from fossils found in the Grand River Canyon (now [[Colorado River]]) of western [[Colorado]], in the [[United States]]. Riggs named the dinosaur ''Brachiosaurus altithorax'', declaring it "the largest known dinosaur". ''Brachiosaurus''  had a proportionally long neck, small skull, and large overall size, all of which are typical for sauropods. Also, the limbs were pillar-like, and the hands columnar. However, the proportions of ''Brachiosaurus'' are unlike most sauropods. The forelimbs were longer than the hindlimbs, which result in a steeply inclined trunk, making the overall body shape reminiscent of a modern [[giraffe]]. Also, while the tail is a typical long dinosaur tail, it was relatively short for a sauropod.
 Much of what is known by laypeople about ''Brachiosaurus'' is in fact based on ''[[Giraffatitan|Giraffatitan brancai]]'', a species of brachiosaurid dinosaur from the [[Tendaguru]] Formation of [[Tanzania]] described by German paleontologist [[Werner Janensch]] as a species of ''Brachiosaurus''. Recent research shows that the differences between the type species of ''Brachiosaurus'' and the Tendaguru material are significant enough that the African material should be placed in a separate genus.
-==Description and paleobiology==
+==Description==
 [[Image:Brachiosaurus DB.jpg|thumb|left|Life restoration]]
 Like all sauropod dinosaurs, ''Brachiosaurus'' was a quadrupedal animal with a small skull, a long neck, a large trunk with a high-ellipsoid cross section, a long, muscular tail and slender, columnar limbs.<ref name=upchurch&al2004>Upchurch, P., Barrett, P.M. & Dodson, P. (2004): "Sauropoda." Pp. 259-322 in Weishampel, D.B., Dodson, P. and Osmolska, H. (eds.): ''The Dinosauria, Second Edition''. University of California Press, Berkeley. ISBN 9780520242098</ref> The skull had a robust, wide muzzle and thick jaw bones, with spoon–shaped teeth.  As in ''Giraffatitan'', there was an arch of bone over the snout and in front of the eyes that encircled the nasal opening, although this arch was not as large as in its relative.<ref name=foster2007b>Foster, J. (2007). "''Brachiosaurus altithorax''." ''Jurassic West: The Dinosaurs of the Morrison Formation and Their World''. Indiana University Press. pp. 205–208.</ref> Large air sacs connected to the lung system were present in the neck and trunk, invading the [[vertebra]]e and ribs, greatly reducing the overall density.<ref name=wedel2003a>Wedel, M.J. (2003). "Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs." ''Paleobiology'' '''29''':243-255.</ref><ref name=wedel2003b>Wedel, M.J. (2003). "The evolution of vertebral pneumaticity in sauropod dinosaurs." ''Journal of Vertebrate Paleontology'' '''23''':344-357.</ref> Unusually for a sauropod, the forelimbs were longer than the hind limbs. The [[humerus]] (upper arm bone) of ''Brachiosaurus'' was relatively lightly built for its size,<ref name=taylor2009>Taylor, M.P. (2009). "[http://www.miketaylor.org.uk/dino/pubs/taylor2009/Taylor2009-brachiosaurus-and-giraffatitan.pdf A re-evaluation of ''Brachiosaurus altithorax'' Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from ''Giraffatitan brancai'' (Janensh 1914)]." ''Journal of Vertebrate Paleontology'', '''29'''(3): 787-806.</ref> measuring {{convert|2.04|m|ft}} in length in the type specimen.<ref name=riggs1903>Riggs, E.S. (1903). [http://books.google.com/books?id=1PjRAAAAMAAJ&pg=PR6&lpg=PR6&dq=%22Brachiosaurus+altithorax,+the+largest+known+dinosaur%22&source=bl&ots=TKCG6wx3LZ&sig=2NQ81hDw0TrzaGD3b6Qi7WmQSCw&hl=en&ei=qr3oS9BUpfA0wOKo1Qk&sa=X&oi=book_result&ct=result&resnum=3&ved=0CBkQ6AEwAg#v=onepage&q=%22Brachiosaurus%20altithorax%2C%20the%20largest%20known%20dinosaur%22&f=false "''Brachiosaurus altithorax'', the largest known dinosaur."] ''American Journal of Science'', '''4'''(15): 299-306.</ref> The [[femur]] (thigh bone) of the type specimen was only {{convert|2.03|m|ft}} long.<ref name=riggs1903/> Unlike other sauropods, ''Brachiosaurus'' appears to have been slightly sprawled at the shoulder joint,<ref name=taylor2009/> and the ribcage was unusually deep.<ref name=riggs1903/> This led to the trunk being inclined, with the front much higher than the hips, and the neck exiting the trunk at a steep angle. Overall, this shape resembles a giraffe more than any other living animal.<ref name=paul1988>{{Cite journal|last=Paul |first=G.S. |year=1988 |title=The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, ''Giraffatitan'', and a comparison of the world's largest dinosaurs |journal=Hunteria |volume=2 |issue=3}}</ref>
@@ Line 44: / Line 44: @@
 Because "Brachiosaurus" ''brancai'' (''Giraffatitan'') is known from much more complete material than ''B. altithorax'', most size estimates for ''Brachiosaurus'' are actually for the African form. There is an additional element of uncertainty for North American ''Brachiosaurus'' because the most complete skeleton appears to have come from a [[subadult]].<ref name=taylor2009/> Over the years, the mass of ''B. altithorax'' has been estimated as {{convert|35.0|MT|ST}},<ref name=paul1988/> {{convert|43.9|MT|ST}},<ref name=Foster2003>{{Cite book|last=Foster |first=J.R. |title=Paleoecological analysis of the vertebrate fauna of the Morrison Formation (Upper Jurassic), Rocky Mountain region, U.S.A. |year=2003 |series=New Mexico Museum of Natural History and Science Bulletin, '''23''' |publisher=New Mexico Museum of Natural History and Science |location=Albuquerque, New Mexico}}</ref> and, most recently, {{convert|28.7|MT|ST}}.<ref name=taylor2009/>  In the first and last cases, the authors also provided estimates for ''Giraffatitan'', and found that genus to be somewhat lighter ({{convert|31.5|MT|ST}} for Paul [1988]<ref name=paul1988/> and {{convert|23.3|MT|ST}} for Taylor [2009]<ref name=taylor2009/>).   The length of ''Brachiosaurus'' has been estimated at {{convert|26|m|ft}}.<ref name=holtz2008>Holtz, T.R. Jr. (2008) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages'' [http://www.geol.umd.edu/~tholtz/dinoappendix/DinoappendixSummer2008.pdf Supplementary Information]</ref>
-===Neck position===
+==Classification==
+''Brachiosaurus'' is the namesake genus of [[Brachiosauridae]].<ref name=riggs1904>{{Cite journal|last=Riggs |first=E.S. |year=1904 |title=Structure and relationships of opisthocoelian dinosaurs. Part II. The Brachiosauridae |journal= Geological Series (Field Columbian Museum) |volume=2 |issue=6 |pages=229–247 |url=http://books.google.com/?id=9y2PAAAAMAAJ&pg=PA246-IA49&lpg=PA246-IA49&dq=%22Structure+and+relationships+of+opisthocoelian+dinosaurs%22#v=onepage&q=%22Structure%20and%20relationships%20of%20opisthocoelian%20dinosaurs%22&f=false}}</ref> Over the years a number of  sauropods have been assigned to Brachiosauridae, such as ''[[Astrodon]]'', ''[[Bothriospondylus]]'', ''[[Dinodocus]]'', ''[[Pelorosaurus]]'', ''[[Pleurocoelus]]'', and ''[[Ultrasaurus]]'',<ref name=lambert1990>{{Cite book|last=Lambert |first=David |coauthors=and the Diagram Group |title=The Dinosaur Data Book |year=1990 |publisher=Avon Books |location=New York |isbn=0-380-75896-3 |chapter=Brachiosaurids |page=142}}</ref> but most of these are currently regarded as [[nomen dubium|dubious]] or of uncertain placement.<ref name=upchurch&al2004/> A [[cladistics|phylogenetic analysis]] of sauropods published in the description of ''[[Abydosaurus]]'' found that genus to form a [[clade]] with ''Brachiosaurus'' and ''Giraffatitan'' (included in ''Brachiosaurus'').<ref name=chure&al2010>{{Cite journal|last=Chure |first=D. |coauthors=Britt, B.; Whitlock, J. A.; and Wilson, J. A. |year=2010 |title=First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition |journal=[[Naturwissenschaften]] |volume=97|pmid=20179896 |issue=4 |pages=379–391|pmc=2841758 |url=http://www.springerlink.com/content/lpn30h8tx2231223/fulltext.pdf |doi=10.1007/s00114-010-0650-6 }}</ref> A more recent analysis focused on possible Asian brachiosaurid material found a clade including ''Abydosaurus'', ''Brachiosaurus'', ''[[Cedarosaurus]]'', ''Giraffatitan'', and ''Paluxysaurus'', but not ''[[Qiaowanlong]]'', the putative Asian brachiosaurid.<ref name=ksepkanorell2010>{{cite journal |last=Ksepka |first=D. T. |coauthors=and Norell, M. A. |year=2010 |title=The illusory evidence for Asian Brachiosauridae: new material of ''Erketu ellisoni'' and a phylogenetic appraisal of basal Titanosauriformes |journal=American Museum Novitates |volume=3700 |pages=1–27 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/6087/1/N3700.pdf |format=pdf}}</ref> Related genera include ''Lusotitan'' and ''[[Sauroposeidon]]''.<ref name=upchurch&al2004/> Brachiosauridae is situated at the base of [[Titanosauriformes]], a group of sauropods that also includes the [[titanosaur]]s.<ref name=ksepkanorell2010/>
-In contrast to most other sauropods, brachiosaurids had an inclined back, due to their long forelimbs. Therefore, if the neck exited the body in a straight line, it already pointed upwards.<ref name=taylor2009/><ref name=stevensparrish1999>Stevens, K. A. and Parrish, M. J. (1999). "Neck posture and feeding habits of two Jurassic sauropod dinosaurs." ''Science'' '''284''':798–800. doi:10.1126/science.284.5415.798</ref><ref name=stevensparrish2005>Stevens, K. A. and Parrish, M. J. (2005). "Digital reconstructions of sauropod dinosaurs and implications for feeding." In ''The sauropods: evolution and paleobiology'' (eds. J. A.Wilson & K. Curry-Rogers), pp. 178–200. Berkeley, CA: University of California Press.</ref><ref name=stevensparrish2005b>Stevens, K. A. and Parrish, M. J. (2005). "Neck posture, dentition and feeding strategies in Jurassic sauropod dinosaurs." In ''Thunder Lizards: The Sauropodomorph dinosaurs" (eds. V. Tidwell & K. Carpenter). Bloomington, IN: Indiana University Press.</ref> The exact angle is influenced by how the pectoral girdle is reconstructed, that is how the shoulder blades are placed on the ribcage.<ref name=stevensparrish1999/><ref name=stevensparrish2005/><ref name=stevensparrish2005b/> The mobility of the neck was reconstructed as quite low by Stevens and Parrish,<ref name=stevensparrish1999/><ref name=stevensparrish2005/><ref name=stevensparrish2005b/> while other researchers like Paul and Christian and Dzemski argued for more flexible necks.<ref name=paul1988/><ref name=dzemskichristian2007>Dzemski, G. and Christian, A. (2007). "Flexibility along the neck of the ostrich (''Struthio camelus'') and consequences for the reconstruction of dinosaurs with extreme neck length." ''Journal of Morphology'' '''268''':701-714.</ref>
+According to the revised diagnosis by Taylor ''Brachiosaurus altithorax'' is diagnosed by a plethora of characters, many to be found on the dorsal (back) vertebrae.<ref name=taylor2009/> Among the characters playing it in the family Brachiosauridae are a ratio of humerus to femur length greater or equal to 0.9 (i.e., the upper arm bone is at least nearly as long as the thigh bone), and a very flattened femur shaft (ratio ≥1.85).<ref name=taylor2009/>
-===Feeding ecology===
-''Brachiosaurus'' is thought to have been a high [[browsing (predation)|browser]], feeding on foliage well above the ground.  Even if it did not hold its neck near vertical, and instead had a straighter neck, its head height may still have been over {{convert|9|m|ft}} above the ground.<ref name=Foster2003/><ref name=foster2007b/> It probably fed mostly on foliage above {{convert|5|m|ft}}. This does not preclude the possibility that it also fed lower at times, between {{convert|3|to|5|m|ft}} up.<ref name=foster2007b/> Its diet likely consisted of ginkgoes, conifers, tree ferns, and large cycads, with intake estimated at {{convert|200|to|400|kg|lb}} of plant matter daily.<ref name=foster2007b/> However, more recent studies estimate that ~{{convert|240|kg|lb}} of plant matter would have been sufficient to feed a {{convert|70|MT|ST}} sauropod,<ref name=hummel&al2008>Hummel, J., Gee, C.T., Südekum, K.-H., Sander, P.M., Nogge, G. and Clauss, M. (2008). "In vitro digestibility of fern and gymnosperm foliage: implications for sauropod feeding ecology and diet selection". ''Proceedings of the Royal Society B'', '''275''':1015-1021. doi:10.1098/rspb.2007.1728</ref> so ''Brachiosaurus'' may have required only about {{convert|120|kg|lb}} of fodder a day. Brachiosaur feeding involved simple up–and–down jaw motion. The teeth were arranged to shear material as they closed, and were probably used to crop and/or nip vegetation.<ref name=bar&up05>{{Cite book|last=Barrett |first=Paul M. |coauthors=and Upchurch, Paul |year=2005 |chapter=Sauropodomorph diversity through time |title=The Sauropods: Evolution and Paleobiology |editors=Curry Rogers, Kristina A.; and Wilson, Jeffrey A. |pages=125–156 |publisher=University of California |location=Berkeley, CA |isbn=0-520-24623-3}}</ref>
-It has repeatedly been suggested, e.g. in the movie ''[[Jurassic Park (film)|Jurassic Park]]'', that ''Brachiosaurus'' could rear into a bipedal or tripodal (with tail support) pose to feed. However, a detailed physical modelling-based analysis of sauropod rearing capabilities by Heinrich Mallison showed that while many sauropods could rear, the unusual brachiosaurid body shape and limb length ratio made them exceptionally ill suited for rearing. The forward position of the center of mass would have led to problems with stability, and required unreasonably large forces in the hips to obtain an upright posture. ''Brachiosaurus'' would also have gained relatively little from rearing (only 33% more feeding height), compared to other sauropods, for which a bipedal pose may have tripled the feeding height.<ref name=mallisonsaubook2>Mallison, H. (in press). "Rearing Giants – kinetic-dynamic modeling of sauropod bipedal and tripodal poses." In Klein, N., Remes, K., Gee, C. & Sander M. (eds): ''Biology of the Sauropod Dinosaurs: Understanding the life of giants.'' Life of the Past (series ed. Farlow, J.). Bloomington, IN: Indiana University Press.</ref>
-===Paleoecology===
-With the removal of the East African ''Giraffatitan'', ''Brachiosaurus'' is known only from the [[Morrison Formation]] of western North America.<ref name=taylor2009/>  The Morrison Formation is interpreted as a [[semiarid]] environment with distinct [[wet season|wet]] and [[dry season]]s,<ref name=russell1989>{{Cite book|last=Russell |first=D. A. |authorlink=Dale Russell |title=An Odyssey in Time: Dinosaurs of North America |year=1989 |publisher=NorthWord  Press |location=Minocqua, Wisconsin |isbn=1-55971-038-1 |pages=64–70 }}</ref><ref name=engelmann&al04>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." ''Sedimentary Geology'' '''167'''(3-4):297-308.</ref> and flat [[floodplain]]s.<ref name=russell1989/> Vegetation varied from [[gallery forest]]s (river–lining forests in otherwise treeless settings) of [[conifer]]s, [[tree fern]]s, and [[fern]]s, to fern [[savanna]]s with rare trees.<ref name=carpenter2006>{{Cite book|last=Carpenter |first=K. |authorlink=Kenneth Carpenter |year=2006  |chapter=Biggest of the big: a critical re-evaluation  of the mega-sauropod  ''Amphicoelias fragillimus'' |editors=Foster, J. R.; and Lucas, S. G. (eds.) |title=Paleontology and Geology of the Upper Jurassic Morrison Formation |series=New Mexico Museum of Natural History and Science Bulletin, '''36'''  |publisher=New Mexico Museum of Natural History and Science |location=Albuquerque, New Mexico |pages=131–138 }}</ref>  Several other sauropod genera were present in the Morrison Formation, with differing body proportions and feeding adaptations.<ref name=Foster2003/> Among these were ''[[Apatosaurus]]'', ''[[Barosaurus]]'', ''[[Camarasaurus]]'', ''[[Diplodocus]]'', ''[[Haplocanthosaurus]]'', and ''[[Supersaurus]]''.<ref name=Foster2003/><ref name=Dchure&al2006>{{Cite book|last=Chure |first=D.J. |coauthors=Litwin, R.; Hasiotis, S.T.; Evanoff, E.; and Carpenter, K. |year=2006 |chapter=The fauna and flora of the Morrison Formation: 2006 |editor=Foster, J.R.; and Lucas, S.G. (eds.) |title=Paleontology and Geology of the Upper Jurassic Morrison Formation |series=New Mexico Museum of Natural History and Science Bulletin, '''36''' |publisher=New Mexico Museum of Natural History and Science |location=Albuquerque, New Mexico |pages=233–248}}</ref> ''Brachiosaurus'' was one of the less abundant Morrison Formation sauropods.  In a survey of over 200 fossil localities, John Foster reported 12 specimens of the genus, comparable to ''Barosaurus'' (13) and ''Haplocanthosaurus'' (12), but far fewer than ''Apatosaurus'' (112), ''Camarasaurus'' (179), and ''Diplodocus'' (98).<ref name=Foster2003/>
-===Metabolism===
-Like all sauropods, ''Brachiosaurus'' was [[warm-blooded|homeothermic]] (maintaining a stable internal temperature) and [[endothermic]] (controlling body temperature through internal means), meaning that it was able to actively control its body temperature, producing the necessary heat through a high [[basal metabolic rate|basic metabolic rate]] of its cells.<ref name=sander&al2010>Sander, P.M., Christian, A., Clauss, M., Fechner, R., Gee, C.T., Griebeler, E.-M., Gunga, H.-C., Hummel, J., Mallison, H., Perry, S.F., Preuschoft, H., Rauhut, O.W.M., Remes, K., Tütken, T., Wings, O. & Witzel, U. (2010). Biology of the sauropod dinosaurs: the evolution of gigantism. ''Biology Reviews'' online first publication, doi:10.1111/j.1469-185X.2010.00137.x http://www3.interscience.wiley.com/cgi-bin/fulltext/123397084/HTMLSTART</ref>
-In the past, ''Brachiosaurus'' has been used an example of a dinosaur for which endothermy is unlikely, because of the combination of great size (leading to overheating) and great [[calorie|caloric]] needs to fuel endothermy.<ref name=russell1989b>{{Cite book|last=Russell |first=D. A. |authorlink=Dale Russell |title=An Odyssey in Time: Dinosaurs of North America |year=1989 |publisher=NorthWord  Press |location=Minocqua, Wisconsin |isbn=1-55971-038-1 |page=78 }}</ref> However, these calculations were based on incorrect assumptions about the available cooling surfaces (the large air sacs were not known), and a grossly inflated body mass. These inaccuracies resulted in overestimation of heat production and underestimation of heat loss.<ref name=sander&al2010/> The large nasal arch has been postulated as an adaptation for cooling the brain, as a surface for evaporative cooling of the blood.<ref name=russell1989b/> Again, when this was proposed, cooling via the air sacs was not known, and thus not taken into account. Furthermore, other similar sized sauropods had no comparable structure. Additionally, in proportion to the entire animal, the nasal arch is very small, and would thus have made only an insignificant contribution to heat loss.
 ==Discovery and history==
 [[Image:Brach humerus.jpg|thumb|Elmer S. Riggs’ assistant lying by a ''Brachiosaurus altithorax'' humerus during the excavation in 1900]]
-The genus ''Brachiosaurus'', and [[type species]] ''B. altithorax'', are based on a partial [[postcrania]]l skeleton from [[Fruita, Colorado|Fruita]], in the valley of the [[Colorado River]] of western Colorado.<ref name=glut1997>{{Cite book|title=Dinosaurs: The Encyclopedia|last=Glut|first=D.F.|publisher=McFarland & Company|year=1997|isbn=0-89950-917-7|chapter=Brachiosaurus|pages=213–221}}</ref>  This specimen was collected from rocks of the Brushy Basin Member of the Morrison Formation<ref name=turner&peterson1999>{{Cite book|last=Turner |first=C.E. |coauthors=and Peterson, F. |year=1999 |chapter=Biostratigraphy of dinosaurs in the Upper Jurassic Morrison Formation of the Western Interior, USA |editor=Gillete, David D. (ed.) |title=Vertebrate Paleontology in Utah |publisher=Utah Geological Survey |location=Salt Lake City, Utah |series=Miscellaneous Publication 99-1 |pages=77–114 |isbn=1-55791-634-9 }}</ref> in [[1900 in paleontology|1900]] by [[Elmer S. Riggs]] and his crew from the Field Columbian Museum (now the [[Field Museum of Natural History]]) of [[Chicago]].<ref name=riggs1903/> It is currently cataloged as [[Field Museum of Natural History|FMNH]]&nbsp;P&nbsp;25107.<ref name=taylor2009/> Riggs and company were working in the area as a result of favorable correspondence between Riggs and S. M. Bradbury, a dentist in nearby [[Grand Junction, Colorado|Grand Junction]].  In 1899 Riggs had sent inquiries to rural locations in the western United States concerning fossil finds, and Bradbury, an amateur collector himself, reported that dinosaur bones had been collected in the area since 1885.<ref name=glut1997/>  It was Riggs' field assistant H. W. Menke who found FMNH&nbsp;P&nbsp;25107,<ref name=riggs1903/> on July 4, 1900, on a small hill later known as Riggs Hill.<ref name=glut1997/> Riggs published a short report in 1901, noting the unusual length of the humerus compared to the femur and the extreme overall size and the resulting giraffe-like proportions, as well as the lesser development of the tail, but did not publish a name for the new dinosaur.<ref name=riggs1901>{{Cite journal|doi=10.1126/science.13.327.549-a|last=Riggs |first=E.S. |year=1091 |title=The largest known dinosaur |journal=Science |volume=13|issue=327 |pages=549–550|pmid=17801098}}</ref> The titles of Riggs (1901) and (1903) suggested that the specimen was the largest known dinosaur.<ref name=riggs1901/><ref name=riggs1903/> Riggs followed his 1903 publication that named ''Brachiosaurus altithorax''<ref name=riggs1903/> with a more detailed description in a [[monograph]] in 1904.<ref name=riggs1904>{{Cite journal|last=Riggs |first=E.S. |year=1904 |title=Structure and relationships of opisthocoelian dinosaurs. Part II. The Brachiosauridae |journal= Geological Series (Field Columbian Musuem) |volume=2 |issue=6 |pages=229–247 |url=http://books.google.com/?id=9y2PAAAAMAAJ&pg=PA246-IA49&lpg=PA246-IA49&dq=%22Structure+and+relationships+of+opisthocoelian+dinosaurs%22#v=onepage&q=%22Structure%20and%20relationships%20of%20opisthocoelian%20dinosaurs%22&f=false}}</ref>
+The genus ''Brachiosaurus'', and [[type species]] ''B. altithorax'', are based on a partial [[postcrania]]l skeleton from [[Fruita, Colorado|Fruita]], in the valley of the [[Colorado River]] of western Colorado.<ref name=glut1997>{{Cite book|title=Dinosaurs: The Encyclopedia|last=Glut|first=D.F.|publisher=McFarland & Company|year=1997|isbn=0-89950-917-7|chapter=Brachiosaurus|pages=213–221}}</ref>  This specimen was collected from rocks of the Brushy Basin Member of the Morrison Formation<ref name=turner&peterson1999>{{Cite book|last=Turner |first=C.E. |coauthors=and Peterson, F. |year=1999 |chapter=Biostratigraphy of dinosaurs in the Upper Jurassic Morrison Formation of the Western Interior, USA |editor=Gillete, David D. (ed.) |title=Vertebrate Paleontology in Utah |publisher=Utah Geological Survey |location=Salt Lake City, Utah |series=Miscellaneous Publication 99-1 |pages=77–114 |isbn=1-55791-634-9 }}</ref> in [[1900 in paleontology|1900]] by [[Elmer S. Riggs]] and his crew from the Field Columbian Museum (now the [[Field Museum of Natural History]]) of [[Chicago]].<ref name=riggs1903/> It is currently cataloged as [[Field Museum of Natural History|FMNH]]&nbsp;P&nbsp;25107.<ref name=taylor2009/> Riggs and company were working in the area as a result of favorable correspondence between Riggs and S. M. Bradbury, a dentist in nearby [[Grand Junction, Colorado|Grand Junction]].  In 1899 Riggs had sent inquiries to rural locations in the western United States concerning fossil finds, and Bradbury, an amateur collector himself, reported that dinosaur bones had been collected in the area since 1885.<ref name=glut1997/>  It was Riggs' field assistant H. W. Menke who found FMNH&nbsp;P&nbsp;25107,<ref name=riggs1903/> on July 4, 1900, on a small hill later known as Riggs Hill.<ref name=glut1997/> Riggs published a short report in 1901, noting the unusual length of the humerus compared to the femur and the extreme overall size and the resulting giraffe-like proportions, as well as the lesser development of the tail, but did not publish a name for the new dinosaur.<ref name=riggs1901>{{Cite journal|doi=10.1126/science.13.327.549-a|last=Riggs |first=E.S. |year=1091 |title=The largest known dinosaur |journal=Science |volume=13|issue=327 |pages=549–550|pmid=17801098}}</ref> The titles of Riggs (1901) and (1903) suggested that the specimen was the largest known dinosaur.<ref name=riggs1901/><ref name=riggs1903/> Riggs followed his 1903 publication that named ''Brachiosaurus altithorax''<ref name=riggs1903/> with a more detailed description in a [[monograph]] in 1904.<ref name=riggs1904/>
 [[Image:Brachiosaurus skull.jpg|thumb|left|The Felch Quarry skull as reconstructed by Carpenter and Tidwell]]
 The Fruita skeleton was not the first discovery of ''Brachiosaurus'' bones, although it was the first to be recognized as belonging to a new and distinct animal.  In 1883, a sauropod skull was found near [[Garden Park, Colorado|Garden Park]], Colorado, at Felch Quarry 1, and was sent to [[Othniel Charles Marsh]] (of "[[Bone Wars]]" fame).<ref name=taylor2009/> Marsh incorporated the skull into his skeletal restoration of "Brontosaurus" (now ''[[Apatosaurus]]'').<ref name=taylor2009/><ref name=marsh1891>{{Cite journal|last=Marsh |first=O.C. |year=1891 |title=Restoration of ''Triceratops'' |journal=American Journal of Science |volume=41 |issue=244 |pages=339–342 |url=http://marsh.dinodb.com/marsh/Marsh%201891%20-%20Restoration%20of%20Triceratops%20%28and%20Brontosaurus%29.pdf |format=pdf}}</ref> It eventually became part of the collections of the [[National Museum of Natural History]], as USNM&nbsp;5730.<ref name=taylor2009/> In the 1970s, when Jack McIntosh and David Berman were working on the issue of the true skull of ''Apatosaurus'', they reevaluated the Garden Park skull as more similar to ''[[Camarasaurus]]''.<ref name=mcintosh&berman1975>{{Cite journal|last=McIntosh |first=J.S. |coauthors=and Berman, D.S. |year=1975 |title=Description of the palate and lower jaw of the sauropod dinosaur ''Diplodocus'' (Reptilia: Saurischia) with remarks on the nature of the skull of ''Apatosaurus'' |journal=Journal of Paleontology |volume=49 |issue=1 |pages=187–199}}</ref> It was described and recognized as a ''Brachiosaurus'' skull in 1998 by [[Kenneth Carpenter]] and Virginia Tidwell, intermediate in [[morphology (biology)|form]] between ''Camarasaurus'' and ''Giraffatitan brancai'' (then still considered to be ''B. brancai'').<ref name=carp&tid1998>Carpenter, K. and Tidwell, V. (1998). "Preliminary description of a ''Brachiosaurus'' skull from Felch Quarry 1, Garden Park, Colorado." Pp. 69–84 in: Carpenter, K., Chure, D. and Kirkland, J. (eds.), ''The Upper Jurassic Morrison Formation: An Interdisciplinary Study''. ''Modern Geology'', '''23''':1-4.</ref>  Because there are no overlapping parts between this skull and FMNH&nbsp;P&nbsp;25107, it cannot be confidently assigned to a species,<ref name=carp&tid1998/><ref name=taylor2009/> so it is classified as ''Brachiosaurus'' sp.<ref name=carp&tid1998/>
@@ Line 68: / Line 57: @@
 Additional discoveries of ''Brachiosaurus'' material in North America have been uncommon and consist of a handful of bones. Material has been described from Colorado,<ref name=Jjenson1987>{{Cite journal|last=Jensen |first=J.A. |year=1987 |title=New brachiosaur material from the Late Jurassic of Utah and Colorado |journal=The Great Basin Naturalist |volume=47 |issue=4 |pages=592–608}}</ref><ref name=curtice&al1996>Curtice, B., Stadtman, K., and Curtice, L. (1996) "A re-assessment of ''Ultrasauros macintoshi'' (Jensen, 1985)." Pp. 87-95 in M. Morales (ed.), ''The Continental Jurassic: Transactions of the Continental Jurassic Symposium'', Museum of Northern Arizona Bulletin number 60.</ref><ref name=curt&stad2001>{{Cite book|last=Curtice |first=B. |coauthors=and Stadtman, K. |year=2001 |chapter=The demise of ''Dystylosaurus edwini'' and a revision of ''Supersaurus vivianae'' |title=Western Association of Vertebrate Paleontologists and Southwest Paleontological Symposium - Proceedings 2001 |editor=McCord, R.D.; and Boaz, D. (eds.) |series=Mesa Southwest Museum Bulletin |volume=8 |pages=33–40}}</ref><ref name=taylor2009/> Oklahoma,<ref name=Bonnan&wedel2004>{{Cite journal|last=Bonnan |first=M.F. |coauthors=and Wedel, M.J. |year=2004 |title=First occurrence of ''Brachiosaurus'' (Dinosauria, Sauropoda) from the Upper Jurassic Morrison Formation of Oklahoma |journal=PaleoBios |volume=24 |issue=2 |pages=12–21}}</ref><ref name=taylor2009/> Utah,<ref name=Jjenson1987/><ref name=taylor2009/> and Wyoming,<ref name=Foster2003/> <ref name=taylor2009/>and undescribed material has been mentioned from several other sites.<ref name=turner&peterson1999/><ref name=taylor2009/> One of these specimens, a [[scapulocoracoid|shoulder blade]] from [[Dry Mesa Quarry]], Colorado, is one of the specimens at the center of the ''[[Supersaurus]]''/''Ultrasauros'' issue of the 1980s and 1990s.  In 1985, [[James A. Jensen]] described disarticulated sauropod remains from the quarry as belonging to several [[taxon|taxa]], including the new genera ''Supersaurus'' and ''Ultrasaurus'',<ref name=jenson1985>{{Cite journal|last=Jensen |first=J.A. |year=1985 |title=Three new sauropod dinosaurs from the Upper Jurassic of Colorado |journal=The Great Basin Naturalist |volume=45 |issue=4 |pages=697–709}}</ref> the latter renamed ''Ultrasauros'' shortly thereafter because [[Ultrasaurus|another sauropod]] already had the name.<ref name=olshevsky1991>Olshevsky, G. (1991). "A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia." ''Mesozoic Meanderings'' '''2''':1-196</ref> Later study showed that the "ultrasaur" material mostly belonged to ''Supersaurus'', although the shoulder blade did not.  Because the holotype of ''Ultrasauros'', a back vertebra, was one of the specimens that was actually from ''Supersaurus'', the name ''Ultrasauros'' is a synonym of ''Supersaurus''.  The shoulder blade is now assigned to ''Brachiosaurus'', but the species is uncertain.<ref name=curtice&al1996/><ref name=taylor2009/> In addition, the Dry Mesa "ultrasaur" was not as large as had been thought; the dimensions of the shoulder's [[coracoid]] bone indicate that the animal was smaller than Riggs' original specimen of ''Brachiosaurus''.<ref name=taylor2009/>
-==Material==
+===Etymology===
+Riggs derived the genus name from the [[Ancient Greek|Greek]] ''brachion''/βραχιων meaning "arm" and ''sauros''/σαυρος meaning "lizard", because he realized that the length of the arms was unusual for a sauropod.<ref name=riggs1903/> The species epithet "altithorax" was chosen because of the unusually deep and wide chest cavity, from [[Latin]] ''altus'' meaning "deep" and Greek ''thorax''/θώραξ (Latin ''thorax''), meaning "breastplate, cuirass, corslet".<ref name=thoraxdict>[http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dqw%2Frac θώραξ], Henry George Liddell, Robert Scott, ''A Greek-English Lexicon'', on Perseus Digital Library</ref>
+==Material and species==
 ===''Brachiosaurus altithorax''===
 [[Image:Altithorax.jpg|thumb|Riggs and H.W. Menke working on ''Brachiosaurus altithorax'' bones]]
@@ Line 87: / Line 79: @@
 There was ample material referred to ''"B." brancai'' in the collections of the [[Museum für Naturkunde]] Berlin, some of which was destroyed during [[World War II]]. Other material was transferred to other institution throughout Germany, some of which was also destroyed. Additional specimens are likely among the material collected by the [[British Museum of Natural History]]'s Tendaguru expedition.<ref name=maier2003>Maier, G. (2003). ''African dinosaurs unearthed. The Tendaguru Expeditions." Bloomington, IN: Indiana University Press.</ref> Much or all of this material probably belongs to ''Giraffatitan'', although some may represent a new brachiosaurid.<ref name=SVPOW2009-11-18>Taylor, M. (18 Nov. 2009): CT-scanning the Archbishop. Sauropod Vertebrate Picture of the Week (Blog) post at http://svpow.wordpress.com/2009/11/18/ct-scanning-the-archbishop/.</ref>
-==Classification==
-''Brachiosaurus'' is the namesake genus of [[Brachiosauridae]].<ref name=riggs1904/> Over the years a number of  sauropods have been assigned to Brachiosauridae, such as ''[[Astrodon]]'', ''[[Bothriospondylus]]'', ''[[Dinodocus]]'', ''[[Pelorosaurus]]'', ''[[Pleurocoelus]]'', and ''[[Ultrasaurus]]'',<ref name=lambert1990>{{Cite book|last=Lambert |first=David |coauthors=and the Diagram Group |title=The Dinosaur Data Book |year=1990 |publisher=Avon Books |location=New York |isbn=0-380-75896-3 |chapter=Brachiosaurids |page=142}}</ref> but most of these are currently regarded as [[nomen dubium|dubious]] or of uncertain placement.<ref name=upchurch&al2004/> A [[cladistics|phylogenetic analysis]] of sauropods published in the description of ''[[Abydosaurus]]'' found that genus to form a [[clade]] with ''Brachiosaurus'' and ''Giraffatitan'' (included in ''Brachiosaurus'').<ref name=chure&al2010>{{Cite journal|last=Chure |first=D. |coauthors=Britt, B.; Whitlock, J. A.; and Wilson, J. A. |year=2010 |title=First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition |journal=[[Naturwissenschaften]] |volume=97|pmid=20179896 |issue=4 |pages=379–391|pmc=2841758 |url=http://www.springerlink.com/content/lpn30h8tx2231223/fulltext.pdf |doi=10.1007/s00114-010-0650-6 }}</ref> A more recent analysis focused on possible Asian brachiosaurid material found a clade including ''Abydosaurus'', ''Brachiosaurus'', ''[[Cedarosaurus]]'', ''Giraffatitan'', and ''Paluxysaurus'', but not ''[[Qiaowanlong]]'', the putative Asian brachiosaurid.<ref name=ksepkanorell2010>{{cite journal |last=Ksepka |first=D. T. |coauthors=and Norell, M. A. |year=2010 |title=The illusory evidence for Asian Brachiosauridae: new material of ''Erketu ellisoni'' and a phylogenetic appraisal of basal Titanosauriformes |journal=American Museum Novitates |volume=3700 |pages=1–27 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/6087/1/N3700.pdf |format=pdf}}</ref> Related genera include ''Lusotitan'' and ''[[Sauroposeidon]]''.<ref name=upchurch&al2004/> Brachiosauridae is situated at the base of [[Titanosauriformes]], a group of sauropods that also includes the [[titanosaur]]s.<ref name=ksepkanorell2010/>
-According to the revised diagnosis by Taylor ''Brachiosaurus altithorax'' is diagnosed by a plethora of characters, many to be found on the dorsal (back) vertebrae.<ref name=taylor2009/> Among the characters playing it in the family Brachiosauridae are a ratio of humerus to femur length greater or equal to 0.9 (i.e., the upper arm bone is at least nearly as long as the thigh bone), and a very flattened femur shaft (ratio ≥1.85).<ref name=taylor2009/>
-===Etymology===
-Riggs derived the genus name from the [[Ancient Greek|Greek]] ''brachion''/βραχιων meaning "arm" and ''sauros''/σαυρος meaning "lizard", because he realized that the length of the arms was unusual for a sauropod.<ref name=riggs1903/> The species epithet "altithorax" was chosen because of the unusually deep and wide chest cavity, from [[Latin]] ''altus'' meaning "deep" and Greek ''thorax''/θώραξ (Latin ''thorax''), meaning "breastplate, cuirass, corslet".<ref name=thoraxdict>[http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dqw%2Frac θώραξ], Henry George Liddell, Robert Scott, ''A Greek-English Lexicon'', on Perseus Digital Library</ref>
 ===Separation from ''Giraffatitan''===
@@ Line 103: / Line 87: @@
 ''Giraffatitan'' was raised to genus level by Olshevsky without comment.<ref name=olshevsky1991/> A detailed study of all material, including the limb and girdle bones, by Michael Taylor in 2009 found that there are significant differences between ''Brachiosaurus altithorax'' and the Tendaguru material in all elements known from both species. Taylor found 26 distinct osteological (bone-based) characters, a larger difference than that between, e.g., ''[[Diplodocus]]'' and ''[[Barosaurus]]'', and therefore argued that the African material should be placed in its own genus, ''Giraffatitan'', as ''G. brancai''.<ref name=taylor2009/> An important difference between the two genera is the overall body shape, with ''Brachiosaurus'' having a 23% longer dorsal (trunk) vertebrate series and a 20 to 25% longer and also taller tail.<ref name=taylor2009/>
+==Paleoecology==
+With the removal of the East African ''Giraffatitan'', ''Brachiosaurus'' is known only from the [[Morrison Formation]] of western North America.<ref name=taylor2009/>  The Morrison Formation is interpreted as a [[semiarid]] environment with distinct [[wet season|wet]] and [[dry season]]s,<ref name=russell1989>{{Cite book|last=Russell |first=D. A. |authorlink=Dale Russell |title=An Odyssey in Time: Dinosaurs of North America |year=1989 |publisher=NorthWord  Press |location=Minocqua, Wisconsin |isbn=1-55971-038-1 |pages=64–70 }}</ref><ref name=engelmann&al04>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." ''Sedimentary Geology'' '''167'''(3-4):297-308.</ref> and flat [[floodplain]]s.<ref name=russell1989/> Vegetation varied from [[gallery forest]]s (river–lining forests in otherwise treeless settings) of [[conifer]]s, [[tree fern]]s, and [[fern]]s, to fern [[savanna]]s with rare trees.<ref name=carpenter2006>{{Cite book|last=Carpenter |first=K. |authorlink=Kenneth Carpenter |year=2006  |chapter=Biggest of the big: a critical re-evaluation  of the mega-sauropod  ''Amphicoelias fragillimus'' |editors=Foster, J. R.; and Lucas, S. G. (eds.) |title=Paleontology and Geology of the Upper Jurassic Morrison Formation |series=New Mexico Museum of Natural History and Science Bulletin, '''36'''  |publisher=New Mexico Museum of Natural History and Science |location=Albuquerque, New Mexico |pages=131–138 }}</ref>  Several other sauropod genera were present in the Morrison Formation, with differing body proportions and feeding adaptations.<ref name=Foster2003/> Among these were ''[[Apatosaurus]]'', ''[[Barosaurus]]'', ''[[Camarasaurus]]'', ''[[Diplodocus]]'', ''[[Haplocanthosaurus]]'', and ''[[Supersaurus]]''.<ref name=Foster2003/><ref name=Dchure&al2006>{{Cite book|last=Chure |first=D.J. |coauthors=Litwin, R.; Hasiotis, S.T.; Evanoff, E.; and Carpenter, K. |year=2006 |chapter=The fauna and flora of the Morrison Formation: 2006 |editor=Foster, J.R.; and Lucas, S.G. (eds.) |title=Paleontology and Geology of the Upper Jurassic Morrison Formation |series=New Mexico Museum of Natural History and Science Bulletin, '''36''' |publisher=New Mexico Museum of Natural History and Science |location=Albuquerque, New Mexico |pages=233–248}}</ref> ''Brachiosaurus'' was one of the less abundant Morrison Formation sauropods.  In a survey of over 200 fossil localities, John Foster reported 12 specimens of the genus, comparable to ''Barosaurus'' (13) and ''Haplocanthosaurus'' (12), but far fewer than ''Apatosaurus'' (112), ''Camarasaurus'' (179), and ''Diplodocus'' (98).<ref name=Foster2003/>
+==Paleobiology==
+===Neck position===
+In contrast to most other sauropods, brachiosaurids had an inclined back, due to their long forelimbs. Therefore, if the neck exited the body in a straight line, it already pointed upwards.<ref name=taylor2009/><ref name=stevensparrish1999>Stevens, K. A. and Parrish, M. J. (1999). "Neck posture and feeding habits of two Jurassic sauropod dinosaurs." ''Science'' '''284''':798–800. doi:10.1126/science.284.5415.798</ref><ref name=stevensparrish2005>Stevens, K. A. and Parrish, M. J. (2005). "Digital reconstructions of sauropod dinosaurs and implications for feeding." In ''The sauropods: evolution and paleobiology'' (eds. J. A.Wilson & K. Curry-Rogers), pp. 178–200. Berkeley, CA: University of California Press.</ref><ref name=stevensparrish2005b>Stevens, K. A. and Parrish, M. J. (2005). "Neck posture, dentition and feeding strategies in Jurassic sauropod dinosaurs." In ''Thunder Lizards: The Sauropodomorph dinosaurs" (eds. V. Tidwell & K. Carpenter). Bloomington, IN: Indiana University Press.</ref> The exact angle is influenced by how the pectoral girdle is reconstructed, that is how the shoulder blades are placed on the ribcage.<ref name=stevensparrish1999/><ref name=stevensparrish2005/><ref name=stevensparrish2005b/> The mobility of the neck was reconstructed as quite low by Stevens and Parrish,<ref name=stevensparrish1999/><ref name=stevensparrish2005/><ref name=stevensparrish2005b/> while other researchers like Paul and Christian and Dzemski argued for more flexible necks.<ref name=paul1988/><ref name=dzemskichristian2007>Dzemski, G. and Christian, A. (2007). "Flexibility along the neck of the ostrich (''Struthio camelus'') and consequences for the reconstruction of dinosaurs with extreme neck length." ''Journal of Morphology'' '''268''':701-714.</ref>
+===Feeding ecology===
+''Brachiosaurus'' is thought to have been a high [[browsing (predation)|browser]], feeding on foliage well above the ground.  Even if it did not hold its neck near vertical, and instead had a straighter neck, its head height may still have been over {{convert|9|m|ft}} above the ground.<ref name=Foster2003/><ref name=foster2007b/> It probably fed mostly on foliage above {{convert|5|m|ft}}. This does not preclude the possibility that it also fed lower at times, between {{convert|3|to|5|m|ft}} up.<ref name=foster2007b/> Its diet likely consisted of ginkgoes, conifers, tree ferns, and large cycads, with intake estimated at {{convert|200|to|400|kg|lb}} of plant matter daily.<ref name=foster2007b/> However, more recent studies estimate that ~{{convert|240|kg|lb}} of plant matter would have been sufficient to feed a {{convert|70|MT|ST}} sauropod,<ref name=hummel&al2008>Hummel, J., Gee, C.T., Südekum, K.-H., Sander, P.M., Nogge, G. and Clauss, M. (2008). "In vitro digestibility of fern and gymnosperm foliage: implications for sauropod feeding ecology and diet selection". ''Proceedings of the Royal Society B'', '''275''':1015-1021. doi:10.1098/rspb.2007.1728</ref> so ''Brachiosaurus'' may have required only about {{convert|120|kg|lb}} of fodder a day. Brachiosaur feeding involved simple up–and–down jaw motion. The teeth were arranged to shear material as they closed, and were probably used to crop and/or nip vegetation.<ref name=bar&up05>{{Cite book|last=Barrett |first=Paul M. |coauthors=and Upchurch, Paul |year=2005 |chapter=Sauropodomorph diversity through time |title=The Sauropods: Evolution and Paleobiology |editors=Curry Rogers, Kristina A.; and Wilson, Jeffrey A. |pages=125–156 |publisher=University of California |location=Berkeley, CA |isbn=0-520-24623-3}}</ref>
+It has repeatedly been suggested, e.g. in the movie ''[[Jurassic Park (film)|Jurassic Park]]'', that ''Brachiosaurus'' could rear into a bipedal or tripodal (with tail support) pose to feed. However, a detailed physical modelling-based analysis of sauropod rearing capabilities by Heinrich Mallison showed that while many sauropods could rear, the unusual brachiosaurid body shape and limb length ratio made them exceptionally ill suited for rearing. The forward position of the center of mass would have led to problems with stability, and required unreasonably large forces in the hips to obtain an upright posture. ''Brachiosaurus'' would also have gained relatively little from rearing (only 33% more feeding height), compared to other sauropods, for which a bipedal pose may have tripled the feeding height.<ref name=mallisonsaubook2>Mallison, H. (in press). "Rearing Giants – kinetic-dynamic modeling of sauropod bipedal and tripodal poses." In Klein, N., Remes, K., Gee, C. & Sander M. (eds): ''Biology of the Sauropod Dinosaurs: Understanding the life of giants.'' Life of the Past (series ed. Farlow, J.). Bloomington, IN: Indiana University Press.</ref>
+===Metabolism===
+Like all sauropods, ''Brachiosaurus'' was [[warm-blooded|homeothermic]] (maintaining a stable internal temperature) and [[endothermic]] (controlling body temperature through internal means), meaning that it was able to actively control its body temperature, producing the necessary heat through a high [[basal metabolic rate|basic metabolic rate]] of its cells.<ref name=sander&al2010>Sander, P.M., Christian, A., Clauss, M., Fechner, R., Gee, C.T., Griebeler, E.-M., Gunga, H.-C., Hummel, J., Mallison, H., Perry, S.F., Preuschoft, H., Rauhut, O.W.M., Remes, K., Tütken, T., Wings, O. & Witzel, U. (2010). Biology of the sauropod dinosaurs: the evolution of gigantism. ''Biology Reviews'' online first publication, doi:10.1111/j.1469-185X.2010.00137.x http://www3.interscience.wiley.com/cgi-bin/fulltext/123397084/HTMLSTART</ref>
+In the past, ''Brachiosaurus'' has been used an example of a dinosaur for which endothermy is unlikely, because of the combination of great size (leading to overheating) and great [[calorie|caloric]] needs to fuel endothermy.<ref name=russell1989b>{{Cite book|last=Russell |first=D. A. |authorlink=Dale Russell |title=An Odyssey in Time: Dinosaurs of North America |year=1989 |publisher=NorthWord  Press |location=Minocqua, Wisconsin |isbn=1-55971-038-1 |page=78 }}</ref> However, these calculations were based on incorrect assumptions about the available cooling surfaces (the large air sacs were not known), and a grossly inflated body mass. These inaccuracies resulted in overestimation of heat production and underestimation of heat loss.<ref name=sander&al2010/> The large nasal arch has been postulated as an adaptation for cooling the brain, as a surface for evaporative cooling of the blood.<ref name=russell1989b/> Again, when this was proposed, cooling via the air sacs was not known, and thus not taken into account. Furthermore, other similar sized sauropods had no comparable structure. Additionally, in proportion to the entire animal, the nasal arch is very small, and would thus have made only an insignificant contribution to heat loss.
 ==In culture==