Brachiosaurus (/ˌbrækiəˈsɔːrəs/) is agenus ofsauropoddinosaur that lived inNorth America during theLate Jurassic, about 155.6 to 145.5 million years ago. It was firstdescribed by American paleontologistElmer S. Riggs in1903 from fossils found in theColorado River valley in westernColorado, United States. Riggs named the dinosaurBrachiosaurus altithorax; thegeneric name isGreek for "arm lizard", in reference to its proportionately long arms, and thespecific name means "deep chest".Brachiosaurus is estimated to have been between 18 and 22 meters (59 and 72 ft) long; body mass estimates of the subadult holotype specimen range from 28.3 to 46.9 metric tons (31.2 to 51.7 short tons). It had a disproportionately long neck, small skull, and large overall size, all of which are typical for sauropods. Atypically,Brachiosaurus had longer forelimbs than hindlimbs, which resulted in a steeply inclinedtrunk, and a proportionally shorter tail.
Brachiosaurus is the namesake genus of thefamilyBrachiosauridae, which includes a handful of other similarsauropods. Most popular depictions ofBrachiosaurus are in fact based onGiraffatitan, a genus of brachiosaurid dinosaur from theTendaguru Formation ofTanzania.Giraffatitan was originally described by German paleontologistWerner Janensch in 1914 as a species ofBrachiosaurus,B. brancai, but moved to its own genus in 2009. Three other species ofBrachiosaurus have been named based onfossils found inAfrica andEurope; two are no longer considered valid, and a third has become a separate genus,Lusotitan.
Thetype specimen ofB. altithorax is still the most complete specimen, and only a few other specimens are thought to belong to the genus, making it one of the rarer sauropods of theMorrison Formation. It is regarded as a highbrowser, possibly cropping or nipping vegetation as high as 9 meters (30 ft) off the ground. Unlike other sauropods, it was unsuited for rearing on its hindlimbs. It has been used as an example of a dinosaur that was most likelyectothermic because of its large size and the corresponding need for sufficientforage, but more recent research suggests it waswarm-blooded. Among the most iconic and initially thought to be one of thelargest dinosaurs,Brachiosaurus has appeared inpopular culture, notably in the 1993 filmJurassic Park.
Riggs and company were working in the area as a result of favorable correspondence between Riggs and Stanton Merill Bradbury, a dentist in nearbyGrand Junction. In the spring of 1899 Riggs had sent letters to mayors in westernColorado, inquiring after possible trails leading from railway heads into northeasternUtah, where he hoped to find fossils ofEocenemammals.[5] To his surprise, he was informed by Bradbury, an amateur collector himself and president of the Western Colorado Academy of Science, that dinosaur bones had been collected near Grand Junction since 1885.[2] Riggs was skeptical of this claim, but his superior, curator of geology Oliver Cummings Farrington, was very eager to add a large sauropod skeleton to the collection to outdo other institutions, and convinced the museum management to invest five hundred dollars in an expedition.[6] Arriving on June 20, 1900, they set camp at the abandoned Goat Ranch.[7] During a prospecting trip on horseback, Riggs's field assistant Harold William Menke found the humerus of FMNHP25107,[4] on July 4,[8] exclaiming it was "the biggest thing yet!". Riggs at first took the find for a badly preservedBrontosaurus specimen and gave priority to excavating Quarry 12, which held a more promisingMorosaurus skeleton. Having secured that, on July 26 he returned to the humerus in Quarry 13, which soon proved to be of enormous size, convincing a puzzled Riggs that he had discovered the largest land animal ever.[9]
An expedition member lying by thehumerus during the excavation in 1900
The site, Riggs Quarry 13, is located on a small hill later known as Riggs Hill; it is today marked by a plaque. MoreBrachiosaurus fossils are reported on Riggs Hill, but other fossil finds on the hill have been vandalized.[8][10] During excavation of the specimen, Riggs misidentified the humerus as a deformed femur due to its great length, and this seemed to be confirmed when an equally-sized, well-preserved real femur of the same skeleton was discovered. In 1904 Riggs noted: "Had it not been for the unusual size of the ribs found associated with it, the specimen would have been discarded as an Apatosaur, too poorly preserved to be of value." It was only after preparation of the fossil material in the laboratory that the bone was recognized as a humerus.[11] The excavation attracted large numbers of visitors, delaying the work and forcing Menke to guard the site to prevent bones from being looted. On August 17, the last bone was jacketed in plaster.[12] After a concluding ten-day prospecting trip, the expedition returned to Grand Junction and hired a team and wagon to transport all fossils to the railway station, during five days; another week was spent to pack them in thirty-eight crates with a weight of 5,700 kilograms (12,500 lb).[13] On September 10, Riggs left for Chicago by train, arriving on the 15th; the railroad companies let both passengers and cargo travel for free, as apublic relations gesture.[14]
The holotype skeleton 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 several ribs.[4][1][15] Riggs described the coracoid as from the left side of the body,[4][11][15] but restudy has shown it to be a right coracoid.[1] At the time of discovery, the lower end of the humerus, the underside of the sacrum, the ilium and the preserved caudal vertebrae were exposed to the air and thus partly damaged by weathering. The vertebrae were only slightly shifted out of their original anatomical position; they were found with their top sides directed downward. The ribs, humerus, and coracoid, however, were displaced to the left side of the vertebral column, indicating transportation by a water current. This is further evidenced by an isolated ilium ofDiplodocus that apparently had drifted against the vertebral column, as well as by a change in composition of the surrounding rocks. While the specimen itself was embedded in fine-grained clay, indicating low-energy conditions at the time of deposition, it was cut off at the seventh presacral vertebra by a thick layer of much coarser sediments consisting of pebbles at its base andsandstone further up, indicating deposition under stronger currents. Based on this evidence, Riggs in 1904 suggested that the missing front part of the skeleton was washed away by a water current, while the hind part was already covered by sediment and thus got preserved.[11]
Riggs (right) and laboratory assistant working on the holotype bones in 1899. The still-jacketedthighbone can be seen on the left.
Riggs published a short report of the new find 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.[15] In 1903, he named thetype speciesBrachiosaurus altithorax.[4] Riggs derived the genus name from theGreekbrachion/βραχίων meaning "arm" andsauros/σαυρος meaning "lizard", because he realized that the length of the arms was unusual for a sauropod.[4] Thespecific epithet was chosen because of the unusually deep and wide chest cavity, fromLatinaltus "deep" and Greekthorax/θώραξ, "breastplate, cuirass, corslet".[16] Latinthorax was derived from the Greek and had become a usual scientific designation for the chest of the body. The titles of Riggs's 1901 and 1903 articles emphasized that the specimen was the "largest-known dinosaur".[4][15] Riggs followed his 1903 publication with a more detailed description in amonograph in 1904.[11]
Preparation of the holotype began in the fall of 1900 shortly after it was collected by Riggs for the Field Museum. First the limb elements were processed. In the winter of 1904, the badly weathered vertebrae of the back and hip were prepared by James B. Abbott and C.T. Kline.[11] As the preparation of each bone was finished, it was put on display in a glass case in Hall 35 of the Fine Arts Palace of theWorlds Columbian Exposition, the Field Museum's first location. All the bones were, solitarily, still on display by 1908 in Hall 35 when the Field Museum's newly mountedApatosaurus was unveiled, the very specimen Riggs had found in Quarry 12,[17] today catalogued as FMNH P25112 and identified as aBrontosaurus exemplar.[18] No mount ofBrachiosaurus was attempted because only twenty percent of the skeleton had been recovered. In 1993, the holotype bones were molded and cast, and the missing bones were sculpted based on material of the relatedBrachiosaurus brancai (nowGiraffatitan) inMuseum für Naturkunde, Berlin. This plastic skeleton was mounted and, in 1994, put on display at the north end of Stanley Field Hall, the main exhibit hall of the Field Museum's current building. The real bones of the holotype were put on exhibit in two large glass cases at either end of the mounted cast. The mount stood until 1999, when it was moved to the BConcourse ofUnited Airlines' Terminal One inO'Hare International Airport to make room for the museum's newly acquiredTyrannosaurus skeleton, "Sue".[19] At the same time, the Field Museum mounted a second plastic cast of the skeleton (designed for outside use) which was on display outside the museum on the NW terrace until 2022.[20][21]
Another outdoor cast was sent toDisney's Animal Kingdom to serve as a gateway icon for the "DinoLand, U.S.A." area, known as the "Oldengate Bridge" that connects the two halves of the fossil quarry themed Boneyard play area.[22]
Assigned material
Composite skeletal reconstruction, scaled to the holotype
Further discoveries ofBrachiosaurus material in North America have been uncommon and consist of a few bones. To date, material can be unambiguously ascribed only to the genus when overlapping with the holotype material, and any referrals of elements from the skull, neck, anterior dorsal region, or distal limbs or feet remain tentative. Nevertheless, material has been described from Colorado,[1][23][24][25] Oklahoma,[1][26] Utah,[1][23] and Wyoming,[1][27] and undescribed material has been mentioned from several other sites.[3][1]
In 1883, farmer Marshall Parker Felch, afossil collector for the American paleontologistOthniel Charles Marsh, reported the discovery of a sauropod skull in Felch Quarry 1, nearGarden Park, Colorado. The skull was found in yellowish white sandstone, near a 1-meter-long (3 ft3+1⁄2 in) cervical vertebra, which was destroyed during an attempt to collect it. The skull was cataloged as YPM 1986, and sent to Marsh at thePeabody Museum of Natural History, who incorporated it into his 1891 skeletal restoration ofBrontosaurus (perhaps because Felch had identified it as belonging to that dinosaur). The Felch Quarry skull consists of the cranium, the maxillae, the right postorbital, part of the left maxilla, the leftsquamosal, the dentaries, and a possible partialpterygoid. The bones were roughly prepared for Marsh, which led to some damage. Felch also collected several postcranial fossils, including a partial cervical vertebra and partial forelimb.[28][29][30] Most of the specimens collected by Felch were sent to theNational Museum of Natural History in 1899 after Marsh's death, including the skull, which was then cataloged as USNM 5730.[31][32][33]
O. C. Marsh's outdated 1891 skeletal reconstruction ofBrontosaurus, with skull inaccurately based on that of the Felch QuarryBrachiosaurus
In 1975 the American paleontologists Jack McIntosh and David Berman investigated the historical issue of whether Marsh had assigned an incorrect skull toBrontosaurus (at the time thought to be ajunior synonym ofApatosaurus), and found the Felch Quarry skull to be of "the generalCamarasaurus type", while suggesting that the vertebra found near it belonged toBrachiosaurus. They concluded that if Marsh had not arbitrarily assigned the Felch quarry skull and anotherCamarasaurus-like skull toBrontosaurus, it would have been recognized earlier that the actual skull ofBrontosaurus andApatosaurus was more similar to that ofDiplodocus.[33] McIntosh later tentatively recognized the Felch Quarry skull as belonging toBrachiosaurus, and brought it to the attention of the American paleontologistsKenneth Carpenter and Virginia Tidwell, while urging them to describe it. They brought the skull to theDenver Museum of Natural History, where they further prepared it and made a reconstruction of it based on casts of the individual bones, with the skulls ofGiraffatitan andCamarasaurus acting as templates for the missing bones.[1][31][34]
In 1998 Carpenter and Tidwell described the Felch Quarry skull, and formally assigned it toBrachiosaurus sp. (of uncertain species), since it is impossible to determine whether it belonged to the speciesB. altithorax itself (as there is no overlapping material between the two specimens). They based the skull's assignment toBrachiosaurus on its similarity to that ofB. brancai, later known asGiraffatitan.[31][34] In 2019, American paleontologists Michael D. D'Emic and Matthew T. Carrano re-examined the Felch Quarry skull after having it further prepared andCT-scanned (while consulting historical illustrations that showed earlier states of the bones), and concluded that aquadrate bone and dentary tooth considered part of the skull by Carpenter and Tidwell did not belong to it. The quadrate is too large to articulate with the squamosal, is preserved differently from the other bones, and was found several meters away. The tooth does not resemble those within the jaws (as revealed by CT data), is larger, and was therefore assigned toCamarasaurus sp. (other teeth assignable to that genus are known from the quarry). They also found it most parsimonious to assign the skull toB. altithorax itself rather than an unspecified species, as there is no evidence of other brachiosaurid taxa in the Morrison Formation (and adding this and other possible elements to aphylogenetic analysis did not change the position ofB. altithorax).[35]
A shoulder blade with coracoid fromDry Mesa Quarry, Colorado, is one of the specimens at the center of theSupersaurus/Ultrasauros issue of the 1980s and 1990s. In 1985James A. Jensen described disarticulated sauropod remains from the quarry as belonging to several exceptionally largetaxa, including the new generaSupersaurus andUltrasaurus,[36] the latter renamedUltrasauros shortly thereafter becauseanother sauropod had already received the name.[37] Later study showed that the "ultrasaur" material mostly belonged toSupersaurus, though the shoulder blade did not. Because the holotype ofUltrasauros, a dorsal vertebra, was one of the specimens that was actually fromSupersaurus, the nameUltrasauros is a synonym ofSupersaurus. The shoulder blade, specimenBYU 9462 (previously BYU 5001), was in 1996 assigned to aBrachiosaurus sp. (of uncertain species) by Brian Curtice and colleagues; in 2009Michael P. Taylor concluded that it could not be referred toB. altithorax.[1][24] 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's original specimen ofBrachiosaurus.[1]
Referred forelimb bone (humerus) from Potter Creek,USNM 21903
Several additional specimens were briefly described by Jensen in 1987.[23] One of these finds, the humerus USNM 21903, was discovered in ca. 1943 by uranium prospectors Vivian and Daniel Jones in the Potter Creek Quarry in western Colorado, and donated to theSmithsonian Institution. Originally, this humerus was part of a poorly preserved partial skeleton that was not collected.[1][23][38] According to Taylor in 2009, it is not clearly referable toBrachiosaurus despite its large size of2.13 meters (6 ft11+3⁄4 in). Jensen himself worked at the Potter Creek site in 1971 and 1975, excavating the disarticulated specimen BYU 4744, which contains a mid-dorsal vertebra, an incomplete left ilium, a left radius and a right metacarpal. According to Taylor in 2009, this specimen can be confidently referred toB. altithorax, as far as it is overlapping with its type specimen. Jensen further mentioned a specimen discovered nearJensen, Utah, that includes a rib2.75 meters (9 ft1⁄4 in) in length, an anterior cervical vertebra, part of a scapula, and a coracoid, although he did not provide a description.[1][23] In 2001, Curtice and Stadtman ascribed two articulated dorsal vertebrae (specimen BYU 13023) from Dry Mesa Quarry toBrachiosaurus.[25] Taylor, in 2009, noted that these vertebrae are markedly shorter than those of theB. altithorax holotype, although otherwise being similar.[1]
In 2012, José Carballido and colleagues reported a nearly complete postcranial skeleton of a small juvenile approximately 2 meters (6 ft 7 in) in length. This specimen, nicknamed "Toni" and cataloged as SMA 0009, stems from the Morrison Formation of theBighorn Basin in north-central Wyoming. Although originally thought to belong to adiplodocid, it was later reinterpreted as a brachiosaurid, probably belonging toB. altithorax.[39] In 2018, the largest sauropod foot ever found was reported from theBlack Hills ofWeston County, Wyoming. The femur is not preserved but comparisons suggest that it was about two percent longer than that of theB. altithorax holotype. Though possibly belonging toBrachiosaurus, the authors cautiously classified it as an indeterminate brachiosaurid.[40] However, the assignment of these two specimens to their respective clades was later questioned by D'Emic and Carrano in 2019. They considered the referral of "Toni" toB. altithorax be based on mistaken interpretations of the species' unique features or of the specimen itself, and deemed it worthy of further study. Analyzing photos of the large foot, D'Emic and Carrano noted that the only feature that allowed referral to Brachiosauridae may have been influenced by damage to the bone it was found on, but did state that "general similarities" withSonorasaurus andGiraffatitan suggested brachiosaurid affinities, but this, the authors stated, would be confirmed only through further study.[28]
Between 1909 and 1912, large-scale paleontological expeditions inGerman East Africa unearthed a considerable amount ofbrachiosaurid material from theTendaguru Formation. In 1914, German paleontologistWerner Janensch listed differences and commonalities between these fossils andB. altithorax, concluding they could be referred to the genusBrachiosaurus. From this material Janensch named two species:Brachiosaurus brancai for the larger and more complete taxon, andBrachiosaurus fraasi for the smaller and more poorly known species.[41] In three further publications in 1929,[42] 1950[43] and 1961,[44] Janensch compared the species in more detail, listing thirteen shared characters betweenBrachiosaurus brancai (which he now considered to includeB. fraasi) andB. altithorax.[1] Taylor, in 2009, considered only four of these characters as valid; six pertain to groups more inclusive than the Brachiosauridae, and the rest are either difficult to assess or refer to material that is notBrachiosaurus.[1]
There was ample material referred toB. brancai in the collections of the Museum für Naturkunde in Berlin, some of which was destroyed duringWorld War II. Other material was transferred to other institutions throughout Germany, some of which was also destroyed. Additional material was collected by theBritish Museum of Natural History's Tendaguru expedition, including a nearly complete skeleton (BMNH R5937) collected by F.W.H. Migeod in 1930. This specimen is now believed torepresent a new species, awaiting description.[1][45]
Janensch based his description ofB. brancai on "Skelett S" (skeleton S) from Tendaguru,[41] but later realized that it comprised two partial individuals: SI and SII.[42] He at first did not designate them as asyntype series, but in 1935 made SI (presently MB.R.2180) thelectotype. Taylor in 2009, unaware of this action, proposed the larger and more complete SII (MB.R.2181) as the lectotype.[1] It includes, among other bones, several dorsal vertebrae, the left scapula, both coracoids, thebreastbones, both humeri, both ulnae and radii (lower arm bones), a right hand, a partial left hand, bothhip bones and the right femur, tibia and fibula (shank bones). Later in 2011, Taylor realized that Janensch had designated the smaller skeleton SI as the lectotype in 1935.[46][47]
Diagram incorporating bones of bothBrachiosaurus andGiraffatitan, byWilliam Diller Matthew, 1915
In 1988Gregory S. Paul published a new reconstruction of the skeleton ofB. brancai, highlighting differences in proportion between it andB. altithorax. Chief among them was a distinction in the way the trunk vertebrae vary: they are fairly uniform in length in the African material, but vary widely inB. altithorax. Paul believed that the limb and girdle elements of both species were very similar, and therefore suggested they be separated not at genus, but only atsubgenus level, asBrachiosaurus (Brachiosaurus) altithorax andBrachiosaurus (Giraffatitan) brancai.[48]Giraffatitan was raised to full genus level by George Olshevsky in 1991, while referring to the vertebral variation.[37] Between 1991 and 2009, the nameGiraffatitan was almost completely disregarded by other researchers.[1]
A detailed 2009 study by Taylor of all material, including the limb and girdle bones, found that there are significant divergences betweenB. altithorax and the Tendaguru material in all elements known from both species. Taylor found twenty-six distinct osteological (bone-based) characters, a larger difference than betweenDiplodocus andBarosaurus, and therefore argued that the African material should indeed be placed in its own genus (Giraffatitan) asGiraffatitan brancai.[1] An important contrast between the two genera is their overall body shape, withBrachiosaurus having a 23 percent longer dorsal vertebral series and a 20 to 25 percent longer and also taller tail.[1] The split was rejected by Daniel Chure in 2010,[49] but from 2012 onward most studies recognized the nameGiraffatitan.[50]
In 1947, at Atalaia in Portugal, brachiosaurid remains were found in layers dating from theTithonian.Albert-Félix de Lapparent and Georges Zbyszewski named them as the speciesBrachiosaurus atalaiensis in 1957.[51] Its referral toBrachiosaurus was doubted in the 2004 edition ofThe Dinosauria by Paul Upchurch, Barret, andPeter Dodson who listed it as an as yet unnamed brachiosaurid genus.[52] Shortly before the publication of the 2004 book, the species had been placed in its own genusLusotitan byMiguel Telles Antunes andOctávio Mateus in 2003.[53] De Lapparent and Zbyszewski had described a series of remains but did not designate atype specimen. Antunes and Mateus selected a partial postcranial skeleton (MIGM4978, 4798, 4801–4810, 4938, 4944, 4950, 4952, 4958, 4964–4966, 4981–4982, 4985, 8807, 8793–87934) as the lectotype; this specimen includes twenty-eight 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 tilting upward indicate thatLusotitan is a brachiosaurid,[53] which was confirmed by some later studies, such as an analysis in 2013.[50]
Brachiosaurus nougaredi
Diagram showing preserved parts of theB. nougaredi sacrum in blue
In 1958 French petroleum geologist F. Nougarède reported to have discovered fragmentary brachiosaurid remains in easternAlgeria, in theSahara Desert.[54] Based on these, Albert-Félix de Lapparent described and named the speciesBrachiosaurus nougaredi in 1960. He indicated the discovery locality as being in theLate Jurassic-age Taouratine Series. He assigned the rocks to this age in part because of the presumed presence ofBrachiosaurus.[55] A more recent review placed it in the "Continental intercalaire", which is considered to belong to theAlbian age of the lateEarly Cretaceous, significantly younger.[52]
The type material moved to Paris consisted of a sacrum, weathered out at the desert surface, and some of the left metacarpals andphalanges. 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 frommetatarsals.[55]
B. nougaredi was in 2004 considered to represent a distinct, unnamed brachiosaurid genus,[52] but a 2013 analysis by Philip D. Mannion and colleagues found that the remains possibly belong to more than one species, as they were collected far apart.[50] The metacarpals were concluded to belong to some indeterminatetitanosauriform. The sacrum was reported lost in 2013. It was not analyzed and provisionally considered to represent an indeterminate sauropod, until such time that it could be relocated in the collections of theMuséum national d'histoire naturelle. Only four out of the five sacralvertebrae are preserved. The total original length was in 1960 estimated at 1.3 meters (4 ft 3 in), compared to 0.91 meters (3 ft 0 in) withB. altithorax.[55] This would make it larger than any other sauropod sacrum ever found, except those ofArgentinosaurus andApatosaurus.[50]
Description
Size compared to a human
Size
Most estimates ofBrachiosaurus altithorax's size are based on the relatedbrachiosauridGiraffatitan (formerly known asB. brancai), which is known from much more complete material thanBrachiosaurus. The two species are the largest brachiosaurids of which relatively extensive remains have been discovered. There is another element of uncertainty for the North AmericanBrachiosaurus because thetype (and most complete) specimen appears to represent asubadult, as indicated by the unfusedsuture between thecoracoid, a bone of theshoulder girdle that forms part of theshoulder joint, and thescapula (shoulder blade).[1] Over the years, the mass of the holotype specimen has been estimated within the range of 28.3–46.9 metric tons (31.2–51.7 short tons).[1][27][48][56][57] Benson et al. suggested a maximum body mass of 56 and 58 metric tons (62 and 64 short tons),[58][59] but these estimates were questioned due to a very large error range and lack of precision.[57] The length ofBrachiosaurus has been estimated at 20–22 meters (66–72ft)[48][60] and 18 meters (59 ft),[56][61] and its height at9.4 meters (30+3⁄4 ft)[61] and 12–13 meters (39–43 ft).[48][62]
While the limb bones of the most completeGiraffatitan skeleton (MB.R.2181) were very similar in size to those of theBrachiosaurus type specimen, the former was somewhat lighter than theBrachiosaurus specimen given its proportional differences. In studies including estimates for both genera,Giraffatitan was estimated at 31.5 metric tons (34.7 short tons),[46][48] 39.5 metric tons (43.5 short tons),[63] 38.0 metric tons (41.9 short tons),[64] 23.3 metric tons (25.7 short tons),[1] and 34.0 metric tons (37.5 short tons).[58] As with the mainBrachiosaurus specimen,Giraffatitan specimen MB.R.2181 likely does not reflect the maximum size of the genus, as afibula (specimenHMXV2) is thirteen percent longer than that of MB.R.2181.[1]
Like all sauropod dinosaurs,Brachiosaurus was a quadruped with a small skull, a long neck, a large trunk with a high-ellipsoid cross section, a long, muscular tail and slender, columnar limbs.[52] Largeair sacs connected to the lung system were present in the neck and trunk, invading thevertebrae and ribs bybone resorption, greatly reducing the overall density of the body.[65][66] The neck is not preserved in theholotype specimen, but was very long even by sauropod standards in the closely relatedGiraffatitan, consisting of thirteen elongatedcervical (neck) vertebrae.[67] The neck was held in a slight S-curve, with the lower and upper sections bent and a straight middle section.[68]Brachiosaurus likely shared withGiraffatitan the very elongatedneck ribs, which ran down the underside of the neck, overlapping several preceding vertebrae. These bony rods were attached to neck muscles at their ends, allowing these muscles to operatedistal portions of the neck while themselves being located closer to the trunk, lightening the distal neck portions.[68][69]
Brachiosaurus andGiraffatitan probably had a small shoulder hump between the third and fifthdorsal (back) vertebra, where the sideward- and upward-directed vertebralprocesses were longer, providing additional surface for neck muscle attachment.[70] The ribcage was deep compared to other sauropods.[4] Though thehumerus (upper arm bone) andfemur (thigh bone) were roughly equal in length, the entire forelimb would have been longer than the hindlimb, as can be inferred from the elongated forearm andmetacarpus of other brachiosaurids.[1] This resulted in an inclined trunk with the shoulder much higher than the hips, and the neck exiting the trunk at a steep angle. The overall build ofBrachiosaurus resembles agiraffe more than any other living animal.[48] In contrast, most other sauropods had a shorter forelimb than hindlimb; the forelimb is especially short in contemporaneousdiplodocoids.[71]
Brachiosaurus differed in its body proportions from the closely relatedGiraffatitan. The trunk was about 25 to 30 percent longer, resulting in a dorsal vertebral column longer than the humerus. Only a single completecaudal (tail) vertebra has been discovered, but its great height suggests that the tail was larger than inGiraffatitan. This vertebra had a much greater area forligament attachment due to a broadenedneural spine, indicating that the tail was also longer than inGiraffatitan, possibly by 20 to 25 percent.[1] In 1988, paleontologist Gregory S. Paul suggested that the neck ofBrachiosaurus was shorter than that ofGiraffatitan, but in 2009, paleontologist Mike P. Taylor pointed out that two cervical vertebrae likely belonging toBrachiosaurus had identical proportions.[1][48] UnlikeGiraffatitan and other sauropods, which had vertically oriented forelimbs, the arms ofBrachiosaurus appear to have been slightly sprawled at the shoulder joints, as indicated by the sideward orientation of the joint surfaces of the coracoids.[1] The humerus was less slender than that ofGiraffatitan, while the femur had similar proportions. This might indicate that the forelimbs ofBrachiosaurus supported a greater fraction of the body weight than is the case forGiraffatitan.[1]
Postcranial skeleton
Vertebral anatomy of the holotype skeleton. Top: sixth dorsal vertebra in back (A) and right side view (B). Bottom: second caudal vertebra in back (C) and side view (D)
Though the vertebral column of the trunk or torso is incompletely known, the back ofBrachiosaurus most likely comprised twelve dorsal vertebrae; this can be inferred from the complete dorsal vertebral column preserved in an unnamed brachiosaurid specimen,BMNH R5937.[72] Vertebrae of the front part of the dorsal column were slightly taller but much longer than those of the back part. This is in contrast toGiraffatitan, where the vertebrae at the front part were much taller but only slightly longer. The centra (vertebral bodies), the lower part of the vertebrae, were more elongated and roughly circular in cross section, while those ofGiraffatitan were broader than tall. Theforamina (small openings) on the sides of the centra, which allowed for the intrusion of air sacs, were larger than inGiraffatitan. The diapophyses (large projections extending sideways from theneural arch of the vertebrae) were horizontal, while those ofGiraffatitan were inclined upward. At their ends, these projections articulated with the ribs; the articular surface was not distinctly triangular as inGiraffatitan. In side view, the upward-projectingneural spines stood vertically and were twice as wide at the base than at the top; those ofGiraffatitan tilted backward and did not broaden at their base. When seen in front or back view, the neural spines widened toward their tops.[1]
InBrachiosaurus, this widening occurred gradually, resulting in a paddle-like shape, while inGiraffatitan the widening occurred abruptly and only in the uppermost portion. At both their front and back sides, the neural spines featured large, triangular and rugose surfaces, which inGiraffatitan were semicircular and much smaller. The various vertebral processes were connected by thin sheets or ridges of bone, which are calledlaminae.Brachiosaurus lacked postspinal laminae, which were present inGiraffatitan, running down the back side of the neural spines. The spinodiapophyseal laminae, which stretched from the neural spines to the diapophyses, were conflated with the spinopostzygapophyseal laminae, which stretched between the neural spines and thearticular processes at the back of the vertebrae, and therefore terminated at mid-height of the neural spines. InGiraffatitan, both laminae were not conflated, and the spinodiapophyseal laminae reached up to the top of the neural spines.Brachiosaurus is further distinguished fromGiraffatitan in lacking three details in the laminae of the dorsal vertebrae that are unique to the latter genus.[1]
Anatomy of the sacrum, ilium, andcoracoid. Top: Sacrum in bottom (A) and right side view (B). Bottom: Right ilium in side view (C) and left coracoid in side view (D)
Air sacs not only invaded the vertebrae but also the ribs. InBrachiosaurus, the air sacs invaded through a small opening on the front side of the rib shafts, while inGiraffatitan openings were present on both the front and back sides of the tuberculum, a bony projection articulating with the diapophyses of the vertebrae. Paul, in 1988, stated that the ribs ofBrachiosaurus were longer than inGiraffatitan, which was questioned by Taylor in 2009.[1] Behind the dorsal vertebral column, thesacrum consisted of five co-ossifiedsacral vertebrae.[11] As inGiraffatitan, the sacrum was proportionally broad and featured very short neural spines. Poor preservation of the sacral material inGiraffatitan precludes detailed comparisons between both genera. Of the tail, only the second caudal vertebra is well preserved.[1]
As inGiraffatitan, this vertebra was slightly amphicoelous (concave on both ends), lacked openings on the sides, and had a short neural spine that was rectangular and tilted backward. In contrast to the second caudal vertebra ofGiraffatitan, that ofBrachiosaurus had a proportionally taller neural arch, making the vertebra about thirty percent taller. The centrum lacked depressions on its sides, in contrast toGiraffatitan. In front or back view, the neural spine broadened toward its tip to approximately three times its minimum width, but no broadening is apparent inGiraffatitan. The neural spines were also inclined backward by about 30 degrees, more than inGiraffatitan (20 degrees). The caudal ribs projected laterally and were not tilted backward as inGiraffatitan. The articular facets of the articular processes at the back of the vertebra were directed downward, while those ofGiraffatitan faced more toward the sides. Besides the articular processes, thehyposphene-hypantrum articulation formed an additional articulation between vertebrae, making the vertebral column more rigid; inBrachiosaurus, the hyposphene was much more pronounced than inGiraffatitan.[1]
Femur (left) and humerus of the holotype
The coracoid was semicircular and taller than broad. Differences fromGiraffatitan are related to its shape in side view, including the straighter suture with the scapula. Moreover, the articular surface that forms part of the shoulder joint was thicker and directed more sideward than inGiraffatitan and other sauropods, possibly indicating a more sprawled forelimb. The humerus, as preserved, measures204 centimeters (80+1⁄2 in) in length, though part of its lower end was lost to erosion; its original length is estimated at 216 centimeters (85 in). This bone was more slender inBrachiosaurus than in most other sauropods, measuring only28.5 centimeters (11+1⁄4 in) in width at its narrowest part. It was, however, more robust than that ofGiraffatitan, being about ten percent broader at the upper and lower ends. At its upper end, it featured a low bulge visible in side view, which is absent inGiraffatitan.[1]
Distinguishing features can also be found in theilium of the pelvis. InBrachiosaurus, the ischiadic peduncle, a downward projecting extension connecting to theischium, reaches farther downward than inGiraffatitan. While the latter genus had a sharp notch between the ischiadic peduncle and the back portion of the ilium, this notch is more rounded inBrachiosaurus. On the upper surface of the hind part of the ilium,Brachiosaurus had a pronounced tubercle that is absent in other sauropods. Of the hindlimb, the femur was very similar to that ofGiraffatitan although slightly more robust, and measured 203 centimeters (80 in) long.[4] As inGiraffatitan, it was strongly elliptical in cross section, being more than twice as wide in front or back view than in side view.[1] Thefourth trochanter, a prominent bulge on the back side of the femoral shaft, was more prominent and located further downward. This bulge served as anchor point for the most important locomotory muscle, thecaudofemoralis, which was situated in the tail and pulled the upper thigh backward when contracted. At the lower end of the femur, the pair ofcondyles did not extend backward as strongly as inGiraffatitan; the two condyles were similar in width inBrachiosaurus but unequal inGiraffatitan.[1]
As reconstructed by Carpenter and Tidwell, the assigned Felch Quarry skull was about 81 centimeters (32 in) long from theoccipital condyle at the back of the skull to the front of thepremaxillae (the front bones of the upper jaw), making it the largest sauropod skull from theMorrison Formation.[31] D'Emic and Carrano instead estimated the skull to have been70 centimeters (27+1⁄2 in) long, and if proportionally similar to that ofGiraffatitan, about55 centimeters (21+1⁄2 in) tall, and 35 centimeters (14 in) wide.[35] Overall, the skull was tall as inGiraffatitan, with a snout that was long (about 36 percent of the skull length according to Carpenter and Tidwell) in front of the nasal bar between the nostrils, typical of brachiosaurids. The snout was somewhat blunt when seen from above (as inGiraffatitan), and since it was set at an angle relative to the rest of the skull, gave the impression of pointing downward.[31][35]
The dorsal andlateral temporal fenestrae (openings at the upper rear and sides of the skull) were large, perhaps due to the force imparted there by the massive jaw adductor musculature. Thefrontal bones on top of the skull were short and wide (similar toGiraffatitan), fused and connected by a suture to theparietal bones, which were also fused together. The surface of the parietals between the dorsal fenestrae was wider than that ofGiraffatitan, but narrower than that ofCamarasaurus. The skull differed from that ofGiraffatitan in its U-shaped (instead of W-shaped) suture between frontal and nasal bones, a shape which appears more pronounced by the frontal bones extending forward over theorbits (eye sockets). Similar toGiraffatitan, the neck of the occipital condyle was very long.[31][35]
Diagram of the Felch Quarry skull, with known material in white
The premaxilla appears to have been longer than that ofCamarasaurus, sloping more gradually toward the nasal bar, which created the very long snout.Brachiosaurus had a long and deepmaxilla (the main bone of the upper jaw), which was thick along the margin where thealveoli (tooth sockets) were placed, thinning upward. Theinterdental plates of the maxilla were thin, fused, porous, and triangular. There were triangularnutrient foramina between the plates, each containing the tip of anerupting tooth. The narialfossa (depression) in front of the bony nostril was long, relatively shallow, and less developed than that ofGiraffatitan. It contained a subnarial fenestra, which was much larger than those ofGiraffatitan andCamarasaurus. The dentaries (the bones of the lower jaws that contained the teeth) were robust, though less than inCamarasaurus. The upper margin of the dentary was arched in profile, but not as much as inCamarasaurus. The interdental plates of the dentary were somewhat oval, with diamond shaped openings between them. The dentary had aMeckelian groove that was open until below the ninth alveolus, continuing thereafter as a shallow trough.[31][35]
Each maxilla had space for fourteen or fifteen teeth, whereasGiraffatitan had eleven andCamarasaurus eight to ten. The maxillae containedreplacement teeth that had rugoseenamel, similar toCamarasaurus, but lacked the smalldenticles (serrations) along the edges. Since the maxilla was wider than that ofCamarasaurus,Brachiosaurus would have had larger teeth. The replacement teeth in the premaxilla had crinkled enamel, and the most complete of these teeth did not have denticles. It was somewhat spatulate (spoon-shaped), and had a longitudinal ridge. Each dentary had space for about fourteen teeth. The maxillary tooth rows ofBrachiosaurus andGiraffatitan ended well in front of theantorbital fenestra (the opening in front of the orbit), whereas they ended just in front of and below the fenestra inCamarasaurus andShunosaurus.[31][35] The teeth ofBrachiosaurus exhibited a high degree of dental complexity relative to other sauropods.[73]
Classification
Replica skeleton outside the FMNH
Riggs, in his preliminary 1903 description of the not yet fully prepared holotype specimen, consideredBrachiosaurus to be an obvious member of the Sauropoda. To determine the validity of the genus, he compared it to the previously named generaCamarasaurus,Apatosaurus,Atlantosaurus, andAmphicoelias, whose validity he questioned given the lack of overlapping fossil material. Because of the uncertain relationships of these genera, little could be said about the relationships ofBrachiosaurus itself.[4] In 1904, Riggs described the holotype material ofBrachiosaurus in more detail, especially the vertebrae. He admitted that he originally had assumed a close affinity withCamarasaurus, but now decided thatBrachiosaurus was more closely related toHaplocanthosaurus. Both genera shared a single line of neural spines on the back and had wide hips. Riggs considered the differences from other taxa significant enough to name a separate family, Brachiosauridae, of whichBrachiosaurus is thenamesake genus. According to Riggs,Haplocanthosaurus was the more primitive genus of the family whileBrachiosaurus was a specialized form.[11]
When describingBrachiosaurus brancai andB. fraasi in 1914, Janensch observed that the unique elongation of the humerus was shared by all threeBrachiosaurus species as well as the BritishPelorosaurus. He also noted this feature inCetiosaurus, where it was not as strongly pronounced as inBrachiosaurus andPelorosaurus.[41] Janensch concluded that the four genera must have been closely related to each other, and in 1929 assigned them to asubfamily Brachiosaurinae within the family Bothrosauropodidae.[42]
During the twentieth century, several sauropods were assigned to Brachiosauridae, includingAstrodon,Bothriospondylus,Pelorosaurus,Pleurocoelus, andUltrasauros.[74] These assignments were often based on broad similarities rather than unambiguoussynapomorphies, shared new traits, and most of these genera are currently regarded asdubious.[52][75] In 1969, in a study by R.F. Kingham,B. altithorax,B. brancai andB. atalaiensis, along with many species now assigned to other genera, were placed in the genusAstrodon, creating anAstrodon altithorax.[76] Kingham's views of brachiosaurid taxonomy have not been accepted by many other authors.[77] Since the 1990s, computer-basedcladistic analyses allow for postulating detailed hypotheses on the relationships between species, by calculating thosetrees that require the fewest evolutionary changes and thus are the most likely to be correct. Such cladistic analyses have cast doubt on the validity of the Brachiosauridae. In 1993, Leonardo Salgado suggested that they were an unnatural group into which all kinds of unrelated sauropods had been combined.[78] In 1997, he published an analysis in which species traditionally considered brachiosaurids were subsequent offshoots of the stem of a larger grouping, theTitanosauriformes, and not a separate branch of their own. This study also pointed out thatB. altithorax andB. brancai did not have any synapomorphies, so that there was no evidence to assume they were particularly closely related.[79]
Fifth dorsal vertebra in front of the pelvis of the holotype, compared to the same region of a human vertebral column
Many cladistic analyses have since suggested that at least some genera can be assigned to the Brachiosauridae, and that this group is a basal branch within the Titanosauriformes.[80] The exact status of each potential brachiosaurid varies from study to study. For example, a 2010 study by Chure and colleagues recognizedAbydosaurus as a brachiosaurid together withBrachiosaurus, which in this study includedB. brancai.[49] In 2009, Taylor noted multiple anatomical differences between the twoBrachiosaurus species, and consequently movedB. brancai into its own genus,Giraffatitan. In contrast to earlier studies, Taylor treated both genera asdistinct units in a cladistic analysis, finding them to besister groups. Another 2010 analysis focusing on possible Asian brachiosaurid material found a clade includingAbydosaurus,Brachiosaurus,Cedarosaurus,Giraffatitan, andPaluxysaurus, but notQiaowanlong, the putative Asian brachiosaurid.[80] Several subsequent analyses have foundBrachiosaurus andGiraffatitan not to be sister groups, but instead located at different positions on the evolutionary tree. A 2012 study by D'Emic placedGiraffatitan in a more basal position, in an earlier branch, thanBrachiosaurus,[77] while a 2013 study by Philip Mannion and colleagues had it the other way around.[50]
Thiscladogram follows that published by Michael D. D'Emic in 2012:[77]
Cladistic analyses also allow scientists to determine which new traits the members of a group have in common, their synapomorphies. According to the 2009 study by Taylor,B. altithorax shares with other brachiosaurids the classic trait of having an upper arm bone that is at least nearly as long as the femur (ratio of humerus length to femur length of at least 0.9). Another shared character is the very flattened femur shaft, its transverse width being at least 1.85 times the width measured from front to rear.[1]
It was believed throughout the nineteenth and early twentieth centuries that sauropods likeBrachiosaurus were too massive to support their own weight on dry land, and instead lived partly submerged in water.[81] Riggs, affirming observations byJohn Bell Hatcher, was the first to defend in length that most sauropods were fully terrestrial animals in his 1904 account onBrachiosaurus, pointing out that their hollow vertebrae have no analogue in living aquatic or semiaquatic animals, and their long limbs and compact feet indicate specialization for terrestrial locomotion.Brachiosaurus would have been better adapted than other sauropods to a fully terrestrial lifestyle through its slender limbs, high chest, wide hips, high ilia and short tail. In its dorsal vertebrae thezygapophyses were very reduced while thehyposphene-hypantrum complex was extremely developed, resulting in a stiff torso incapable of bending sideways. The body was fit for only quadrupedal movement on land.[11] Though Riggs's ideas were gradually forgotten during the first half of the twentieth century, the notion of sauropods as terrestrial animals has gained support since the 1950s, and is now universally accepted among paleontologists.[81][82] In 1990 the paleontologist Stephen Czerkas stated thatBrachiosaurus could have entered water occasionally to cool off (thermoregulate).[83]Brachiosaurus is estimated to have walked at a speed of 1.49 meters per second (5.4 km/h; 3.3 mph).[84]
Ongoing debate revolves around the neck posture of brachiosaurids, with estimates ranging from near-vertical to horizontal orientations.[85] The idea of near-vertical postures in sauropods in general was popular until 1999, when Stevens and Parrish argued that the sauropod neck was not flexible enough to be held in an upright, S-curved pose, and instead was held horizontally.[70][86] Reflecting this research, various newspapers ran stories criticizing the Field MuseumBrachiosaurus mount for having an upward curving neck. Museum paleontologists Olivier Rieppel and Christopher Brochu defended the posture in 1999, noting the long forelimbs and upward sloping backbone. They also stated that the most developed neural spines for muscle attachment being positioned in the region of the shoulder girdle would have permitted the neck to be raised in a giraffe-like posture. Furthermore, such a pose would have required less energy than lowering its neck, and the inter-vertebral discs would not have been able to counter the pressure caused by a lowered head for extended periods of time (though lowering its neck to drink must have been possible).[87] Some recent studies also advocated a more upward directed neck. Christian and Dzemski (2007) estimated that the middle part of the neck inGiraffatitan was inclined by sixty to seventy degrees; a horizontal posture could be maintained only for short periods of time.[68]
With their heads held high above the heart, brachiosaurids would have had stressed cardiovascular systems. It is estimated that the heart ofBrachiosaurus would have to pump double the blood pressure of a giraffe to reach the brain, and possibly weighed 400 kg (880 lb).[88] The distance between head and heart would have been reduced by the S-curvature of the neck by more than2 meters (6+1⁄2 ft) in comparison to a totally vertical posture. The neck may also have been lowered during locomotion by twenty degrees.[68] In studying the inner ear ofGiraffatitan, Gunga & Kirsch (2001) concluded that brachiosaurids would have moved their necks in lateral directions more often than in dorsal-ventral directions while feeding.[68][89]
Feeding and diet
Replica skeleton showing the neck pointing upward, atO'Hare International Airport (formerly housed in the Field Museum)
Brachiosaurus is thought to have been a highbrowser, feeding on foliage well above the ground. Even if it did not hold its neck near vertical, and instead had a less inclined neck, its head height may still have been over 9 meters (30 ft) above the ground.[27][61] It probably fed mostly on foliage above 5 meters (16 ft). This does not preclude the possibility that it also fed lower at times, between 3 and 5 meters (9.8 and 16.4 ft) up.[61] Its diet likely consisted of ginkgos, conifers, tree ferns, and large cycads, with intake estimated at 200 to 400 kilograms (440 to 880 lb) of plant matter daily in a 2007 study.[61] Brachiosaurid feeding involved simple up-and-down jaw motion.[90] As in other sauropods, animals would have swallowed plant matter without further oral processing, and relied onhindgut fermentation for food processing.[85] The teeth were somewhat spoon-shaped and chisel-like.[91] Such teeth are optimized for non-selective nipping,[92] and the relatively broad jaws could crop large amounts of plant material.[91] Even if aBrachiosaurus of forty tonnes would have needed half a tonne of fodder, its dietary needs could have been met by a normal cropping action of the head. If it fed sixteen hours per day, biting off between a tenth and two-thirds of a kilogram, taking between one and six bites per minute, its daily food intake would have equaled roughly 1.5 percent of its body mass, comparable to the requirement of a modern elephant.[93]
AsBrachiosaurus shared itshabitat, the Morrison, with many other sauropod species, its specialization for feeding at greater heights would have been part of a system ofniche partitioning, the various taxa thus avoiding direct competition with each other. A typical food tree might have resembledSequoiadendron. The fact that such tall conifers were relatively rare in the Morrison might explain whyBrachiosaurus was much less common in itsecosystem than the relatedGiraffatitan, which seems to have been one of the most abundant sauropods in the Tendaguru.[94]Brachiosaurus, with its shorter arms and lower shoulders, was not as well-adapted to high-browsing asGiraffatitan.[95]
It has been suggested thatBrachiosaurus could rear on its hind legs to feed, using its tail for extra ground support.[48] A detailed physical modelling-based analysis of sauropod rearing capabilities by Heinrich Mallison showed that while many sauropods could rear, the unusual body shape and limb length ratio of brachiosaurids made them exceptionally ill-suited for rearing. The forward position of its 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 only 33 percent more feeding height, compared to other sauropods, for which rearing may have tripled the feeding height.[96] A bipedal stance might have been adopted byBrachiosaurus in exceptional situations, like male dominance fights.[97]
The downward mobility of the neck ofBrachiosaurus would have allowed it to reach open water at the level of its feet, while standing upright. Modern giraffes spread their forelimbs to lower the mouth in a relatively horizontal position, to more easily gulp down the water. It is unlikely thatBrachiosaurus could have attained a stable posture this way, forcing the animal to plunge the snout almost vertically into the surface of a lake or stream. This would have submerged its fleshy nostrils if they were located at the tip of the snout as Witmer hypothesized. Hallett and Wedel therefore in 2016 rejected his interpretation and suggested that they were in fact placed at the top of the head, above the bony nostrils, as traditionally thought. The nostrils might have evolved their retracted position to allow the animal to breathe while drinking.[98]
Nostril function
The fleshy external nostril would have been placed at the front of the nasal fossa, here demarcated by a dashed line.
The bony nasal openings ofneosauropods likeBrachiosaurus were large and placed on the top of their skulls. Traditionally, the fleshy nostrils of sauropods were thought to have been placed likewise on top of the head, roughly at the rear of the bony nostril opening, because these animals were erroneously thought to have been amphibious, using their large nasal openings as snorkels when submerged. The American paleontologistLawrence M. Witmer rejected this reconstruction in 2001, pointing out that all livingvertebrate land animals have their external fleshy nostrils placed at the front of the bony nostril. The fleshy nostrils of such sauropods would have been placed in an even more forward position, at the front of the narial fossa, the depression which extended far in front of the bony nostril toward the snout tip.[99]
Czerkas speculated on the function of the peculiar brachiosaurid nose, and pointed out that there was no conclusive way to determine where the nostrils where located, unless a head with skin impressions was found. He suggested that the expanded nasal opening would have made room for tissue related to the animal's ability to smell, which would have helped smell proper vegetation. He also noted that in modern reptiles, the presence of bulbous, enlarged, and uplifted nasal bones can be correlated with fleshy horns and knobby protuberances, and thatBrachiosaurus and other sauropods with large noses could have hadornamental nasal crests.[83]
It has been proposed that sauropods, includingBrachiosaurus, may have hadproboscises (trunks) based on the position of the bony narial orifice, to increase their upward reach. Fabien Knoll and colleagues disputed this forDiplodocus andCamarasaurus in 2006, finding that the opening for thefacial nerve in the braincase was small. The facial nerve was thus not enlarged as in elephants, where it is involved in operating the sophisticated musculature of the proboscis. However, Knoll and colleagues also noted that the facial nerve forGiraffatitan was larger, and could therefore not discard the possibility of a proboscis in this genus.[100]
Metabolism
Like other sauropods,Brachiosaurus was probablyhomeothermic (maintaining a stable internal temperature) andendothermic (controlling body temperature through internal means) at least while growing, meaning that it could actively control its body temperature ("warm-blooded"), producing the necessary heat through a highbasic metabolic rate of its cells.[85] Russel (1989) usedBrachiosaurus as an example of a dinosaur for which endothermy is unlikely, because of the combination of great size (leading to overheating) and greatcaloric needs to fuel endothermy.[101] Sander (2010) found that these calculations were based on incorrect body mass estimates and faulty assumptions on the available cooling surfaces, as the presence of large air sacs was unknown at the time of the study. These inaccuracies resulted in the overestimation of heat production and the underestimation of heat loss.[85] The large nasal arch has been postulated as an adaptation for cooling the brain, as a surface forevaporative cooling of the blood.[101]
Air sacs
Dorsal vertebrae of the holotype. The openings at their lower sides arepleurocoels, through whichair sacs invaded the bone and connected with air cells inside.
Therespiration system of sauropods, like that of birds, made use of air sacs. There was not a bidirectional airflow as with mammals, in which the lungs function asbellows, first inhaling and then exhaling air. Instead the air was sucked from thetrachea into an abdominal air sac in the belly which then pumped it forward through the parabronchi, air loops, of the stiff lung. Valves prevented the air from flowing backward when the abdominal air sac filled itself again; at the same time a cervical air sac at the neck base sucked out the spent air from the lung. Both air sacs contracted simultaneously to pump the used air out of the trachea. This procedure guaranteed a unidirectional airflow, the air always moving in a single forward direction in the lung itself. This significantly improved theoxygen intake and the release ofcarbon dioxide. Not only was dead air removed quickly but also the blood flow in the lung was counterdirectional in relation to the airflow, leading to a far more effectivegas exchange.[102]
In sauropods, the air sacs did not simply function as an aid for respiration; by means of air channels they were connected to much of the skeleton. These branches, thediverticula, via pneumatic openings invaded many bones and strongly hollowed them out. It is not entirely clear what the evolutionary benefit of this phenomenon was but in any case it considerably lightened the skeleton. They might also have removed excess heat to aid thermoregulation.[102]
In 2016, Mark Hallett and Mathew Wedel for the first time reconstructed the entire air sac system of a sauropod, usingB. altithorax as an example of how such a structure might have been formed. In their reconstruction a large abdominal air sac was located between the pelvis and the outer lung side. As with birds, three smaller sacs assisted the pumping process from the underside of the breast cavity: at the rear the posterior thoracic air sac, in the middle the anterior thoracic air sac and in front the clavicular air sac, in that order gradually diminishing in size. The cervical air sac was positioned under the shoulder blade, on top of the front lung. The air sacs were via tubes connected with the vertebrae. Diverticula filled the various fossae andpleurocoels that formed depressions in the vertebral bone walls. These were again connected with inflexible air cells inside the bones.[102]
Growth
JuvenileB. sp. specimen SMA 0009, nicknamed Toni. The skull is reconstructed following the initialdiplodocoid identification),Sauriermuseum Aathal.
Theontogeny ofBrachiosaurus has been reconstructed by Carballido and colleagues in 2012 based on Toni (SMA 0009), a postcranial skeleton of a young juvenile with an estimated total body length of just 2 meters (6.6 ft). This skeleton shares some unique traits with theB. altithorax holotype, indicating it is referable to this species. These commonalities include an elevation on the rear blade of the ilium; the lack of a postspinal lamina; vertical neural spines on the back; an ilium with a subtle notch between the appendage for the ischium and the rear blade; and the lack of a side bulge on the upper thighbone. There are also differences; these might indicate that the juvenile is not aB. altithorax individual after all, but belongs to a new species. Alternatively, they might be explained as juvenile traits that would have changed when the animal matured.[103]
Such ontogenetic changes are especially to be expected in the proportions of an organism. The middle neck vertebrae of SMA 0009 are remarkably short for a sauropod, being just 1.8 times longer than high, compared with a ratio of 4.5 inGiraffatitan. This suggests that the necks of brachiosaurids became proportionally much longer while their backs, to the contrary, experienced relative negative growth. The humerus of SMA 0009 is relatively robust: it is more slender than that of most basal titanosauriforms but thicker than the upper arm bone ofB. altithorax. This suggests that it was already lengthening in an early juvenile stage and became even more slender during growth. This is in contrast to diplodocoids and basal macronarians, whose slender humeri are not due to suchallometric growth.Brachiosaurus also appears to have experienced an elongation of the metacarpals, which in juveniles were shorter compared to the length of the radius; SMA 0009 had a ratio of just 0.33, the lowest known in the entire Neosauropoda.[103]
Another plausible ontogenetic change is the increasedpneumatization of the vertebrae. During growth, thediverticula of the air sacs invaded the bones and hollowed them out. SMA 0009 already has pleurocoels, pneumatic excavations, at the sides of its neck vertebrae. These are divided by a ridge but are otherwise still very simple in structure, compared with the extremely complex ridge systems typically shown by adult derived sauropods. Its dorsal vertebrae still completely lack these.[103]
Two traits are not so obviously linked to ontogeny. The neural spines of the rear dorsal vertebrae and the front sacral vertebrae are extremely compressed transversely, being eight times longer from front to rear than wide from side to side. The spinodiapophyseal lamina or "SPOL", the ridge normally running from each side of the neural spine toward each diapophysis, the transverse process bearing the contact facet for the upper rib head, is totally lacking. Both traits could beautapomorphies, unique derived characters proving that SMA 0009 represents a distinct species, but there are indications that these traits are growth-related as well. Of the basal sauropodTazoudasaurus a young juvenile is known that also lacks the spinodiapophyseal lamina, whereas the adult form has an incipient ridge. Furthermore, a very young juvenile ofEuropasaurus had a weak SPOL but it is well developed in mature individuals. These two cases represent the only finds in which the condition can be checked; they suggest that the SPOL developed during growth. As this very ridge widens the neural spine, its transverse compression is not an independent trait and the development of the SPOL plausibly precedes the thickening of the neural spine with more mature animals.[103]
Sauropods were likely able to sexually reproduce before they attained their maximum individual size. The maturation rate differed between species. Its bone structure indicates thatBrachiosaurus was able to reproduce when it reached forty percent of its maximal size.[104] In 2005,Jack Horner estimated that Brachiosaurus reached adulthood in less than 20 years.[105]
Paleoecology
Map showing locations ofbrachiosaurid remains from theMorrison Formation (gray); 5(the red dot) is theB. altithorax type locality
Brachiosaurus is known only from the Morrison Formation of western North America (following the reassignment of the African species), from theKimmeridgian andTithonian ages (155.6 to 145.5 million years ago).[1] The Morrison Formation is interpreted as asemiarid environment with distinctwet anddry seasons,[106][107] and flatfloodplains.[106] Several other sauropod genera were present in the Morrison Formation, with differing body proportions and feeding adaptations.[27][108] Among these wereApatosaurus,Barosaurus,Camarasaurus,Diplodocus,Haplocanthosaurus, andSupersaurus.[27][109]Brachiosaurus was one of the less abundant Morrison Formation sauropods. In a 2003 survey of more than two hundred fossil localities, John Foster reported 12 specimens of the genus, comparable toBarosaurus (13) andHaplocanthosaurus (12), but far fewer thanApatosaurus (112),Camarasaurus (179), andDiplodocus (98).[27] If the large foot reported from Wyoming (the northernmost occurrence of a brachiosaurid in North America) did belong toBrachiosaurus, the genus would have covered a wide range of latitudes.Brachiosaurids could process tough vegetation with their broad-crowned teeth, and might therefore have covered a wider range of vegetational zones than for example diplodocids.Camarasaurids, which were similar in tooth morphology to brachiosaurids, were also widespread and are known to have migrated seasonally, so this might have also been true for brachiosaurids.[40]
Riggs in the first instance tried to limit public awareness of the find. When reading a lecture to the inhabitants of Grand Junction, illustrated by lantern slides, on July 27, 1900, he explained the general evolution of dinosaurs and the exploration methods of museum field crews but did not mention that he had just found a spectacular specimen.[113] He feared that teams of other institutions might soon learn of the discovery and take away the best of the remaining fossils. A week later, his host Bradbury published an article in the localGrand Junction News announcing the find of one of the largest dinosaurs ever. On August 14,The New York Times brought the story.[114] At the time sauropod dinosaurs appealed to the public because of their great size, often exaggerated by sensationalist newspapers.[115] Riggs in his publications played into this by emphasizing the enormous magnitude ofBrachiosaurus.[116] Replica skeletons ofBrachiosaurus can be seen in Chicago, Illinois, one outside the Field Museum and another inside theO'Hare International Airport.[117][118][119][120][121]
Brachiosaurus has been called one of the most iconic dinosaurs, but most popular depictions are based on the African speciesB. brancai which has since been moved to its own genus,Giraffatitan.[1] Amain beltasteroid,1991 GX7, was named9954 Brachiosaurus in honor of the genus in 1991.[122][123]Brachiosaurus was featured in the 1993 movieJurassic Park, as the firstcomputer generated dinosaur shown.[124] These effects were considered ground-breaking at the time, and the awe of the movie's characters upon seeing the dinosaur for the first time was mirrored by audiences.[125][126] The movements of the movie'sBrachiosaurus were based on the gait of a giraffe combined with the mass of an elephant. A scene later in the movie used ananimatronic head and neck, for when aBrachiosaurus interacts with human characters.[124] The digital model ofBrachiosaurus used inJurassic Park later became the starting point for theronto models in the 1997 special edition of the filmStar Wars Episode IV: A New Hope.[127]
^abTurner, C.E.; Peterson, F. (1999). "Biostratigraphy of dinosaurs in the Upper Jurassic Morrison Formation of the Western Interior, USA". In Gillete, David D. (ed.).Vertebrate Paleontology in Utah. Miscellaneous Publication 99-1. Salt Lake City, Utah: Utah Geological Survey. pp. 77–114.ISBN978-1-55791-634-1.
^abChenoweth, W.L. (1987). "The Riggs Hill and Dinosaur Hill sites, Mesa County, Colorado". In Averett, Walter R. (ed.).Paleontology and Geology of the Dinosaur Triangle. Grand Junction, Colorado: Museum of Western Colorado. pp. 97–100.LCCN93247073.OCLC680488874.
^Liddell, H.G.; Scott, R."θώραξ".A Greek-English Lexicon. Perseus Digital Library.Archived from the original on October 19, 2022. RetrievedApril 6, 2018.
^abcdeJensen, J.A. (1987). "New brachiosaur material from the Late Jurassic of Utah and Colorado".The Great Basin Naturalist.47 (4):592–608.
^abCurtice, B.; Stadtman, K.; Curtice, L. (1996). "A re-assessment ofUltrasauros macintoshi (Jensen, 1985)". In Morales, M. (ed.).The Continental Jurassic: Transactions of the Continental Jurassic Symposium. Vol. 60. Museum of Northern Arizona Bulletin. pp. 87–95.
^abCurtice, B.; Stadtman, K. (2001). "The demise ofDystylosaurus edwini and a revision ofSupersaurus vivianae". In McCord, R.D.; Boaz, D. (eds.).Western Association of Vertebrate Paleontologists and Southwest Paleontological Symposium – Proceedings 2001. Vol. 8. Mesa Southwest Museum Bulletin. pp. 33–40.
^abcdefghCarpenter, K.; Tidwell, V. (1998). "Preliminary description of aBrachiosaurus skull from Felch Quarry 1, Garden Park, Colorado".Modern Geology.23 (1–4):69–84.
^abMcIntosh, J.S.; Berman, D.S. (1975). "Description of the palate and lower jaw of the sauropod dinosaurDiplodocus (Reptilia: Saurischia) with remarks on the nature of the skull ofApatosaurus".Journal of Paleontology.49 (1):187–199.JSTOR1303324.
^abTidwell, V. (1996). "Restoring crushed Jurassic dinosaur skulls for display". In Morales, M. (ed.).The Continental Jurassic: Transactions of the Continental Jurassic Symposium. Vol. 60. Museum of Northern Arizona Bulletin.
^abcJanensch, W. (1929). "Material und Formengehalt der Sauropoden in der Ausbeute der Tendaguru-Expedition" [Material and molds of the sauropod yield of the Tendaguru Expedition].Palaeontographica (in German).2 (Suppl. 7):1–34.
^Janensch, W. (1950). "Die Wirbelsäule vonBrachiosaurus brancai" [The spine ofBrachiosaurus brancai].Palaeontographica (in German).3 (Suppl. 7):27–93.
^de Lapparent, A.F.; Zbyszewski, G. (1957)."Les dinosauriens du Portugal"(PDF).Mémoire Service Géologique Portugal.2:1–63. Archived fromthe original(PDF) on January 30, 2022. RetrievedMarch 12, 2018.
^Lapparent, A.F. de; Claracq, P.; Nougarède, F. (1958). "Nouvelles découvertes de Vertébrés dans les séries continentales au Nord d'Edjelch (Sahara central)".Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences de Paris.247:2399–2402.
^abcde Lapparent, A. F. (1960)."Les dinosauriens du "continental intercalaire" du Sahara central"" [The dinosaurs of the "continental intercalaire" of the central Sahara](PDF).Mémoires de la Société Géologique de France. Nouvelle Séries (in French).39 (1–6). Translated by Carrano, Matthew:1–57. Archived fromthe original(PDF) on June 17, 2021. RetrievedSeptember 25, 2010.
^abcdeFoster, J. (2007). "Brachiosaurus altithorax".Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indianapolis: Indiana University Press. pp. 205–208.ISBN978-0-253-34870-8.
^Klein, Nicole; Remes, Kristian; Gee, Carole T.; Sander, P. Martin (2011). "Appendix: Compilation of published body mass data for a variety of basal sauropodomorphs and sauropods".Biology of the sauropod dinosaurs. Indiana University Press. pp. 317–320.ISBN978-0-253-35508-9.
^Wedel, Mathew J.; Cifelli, R. L.; Sanders, R. K. (2000). "Osteology, paleobiology, and relationships of the sauropod dinosaurSauroposeidon".Acta Palaeontologica Polonica.45:343–388.
^Kingham, R.F. (1962). "Studies of the sauropod dinosaurAstrodon Leidy".Proceedings of the Washington Junior Academy of Sciences.1:38–44.
^Salgado, L., R. A. Coria, and J. O. Calvo. 1997. "Evolution of titanosaurid sauropods. I: phylogenetic analysis based on the postcranial evidence".Ameghiniana34: 3-32
^Scott A., Lee; Justyna, Slowiak (2025). "Sauntering Sauropods: The Preferred Walking Speeds of the Largest Land Animals That Ever Lived".The Physics Teacher.63 (1):20–22.Bibcode:2025PhTea..63a..20L.doi:10.1119/5.0187569.
^Fastovsky, D. E.; Weishampel, D. B. (2016).Dinosaurs: A Concise Natural History. Cambridge University Press. p. 206.ISBN978-1-107-13537-6.
^Gunga, H.-C.; Kirsch, K. (2001). "Von Hochleistungsherzen und wackeligen Hälsen" [Of high performance hearts and wobbly necks].Forschung (in German).2–3:4–9.
^Mallison, H. (2011). "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.
^Foster, J. (2007). "Appendix". Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 327–329.
^Chure, D.J.; Litwin, Rl; Hasiotis, S.T.; Evanoff, E.; Carpenter, K. (2006)."The fauna and flora of the Morrison Formation: 2006". In Foster, J.R.; Lucas, S.G. (eds.).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.Archived from the original on September 2, 2017. RetrievedSeptember 12, 2018.
^Chure, Daniel J.; Litwin, Ron; Hasiotis, Stephen T.; Evanoff, Emmett; Carpenter, K. (2006)."The fauna and flora of the Morrison Formation: 2006". In Foster, John R.; Lucas, Spencer G. (eds.).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.Archived from the original on September 2, 2017. RetrievedSeptember 12, 2018.
^Foster, John 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. p. 29.
^Carpenter, K. (2006). "Biggest of the big: a critical re-evaluation of the mega-sauropodAmphicoelias fragillimus". In Foster, John R.; Lucas, Spencer G. (eds.).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.
^"Ronto".Databank. Star Wars.com. Archived fromthe original on October 3, 2008. RetrievedJanuary 13, 2009.
Bibliography
Brinkman, P. D. (2010),The Second Jurassic Dinosaur Rush: Museums and Paleontology in America at the Turn of the Twentieth Century, Chicago and London: The University of Chicago Press,ISBN978-0-226-07472-6
Hallett, M.; Wedel, M. (2016),The Sauropod Dinosaurs: Life in the Age of Giants, Baltimore: Johns Hopkins University Press,ISBN978-1-4214-2028-8
External links
The dictionary definition ofBrachiosaurus at Wiktionary
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