Brachiosaurus

Holotype specimen e, sacrum, ilium and ribs are in view. The genus Brachiosaurus is based on a partial postcranial skeleton discovered in 1900 in the valley of the Colorado River near Fruita, Colorado. This specimen, which was later declared the holotype, comes from rocks of the Brushy Basin Member of the Morrison Formation, and therefore is late Kimmeridgian in age, about 154to 153million years old. Discovered by American paleontologist Elmer S. Riggs and his crew from the Field Columbian Museum (now the Field Museum of Natural History) of Chicago, exclaiming it was "the biggest thing yet!". Riggs at first took the find for a badly preserved Brontosaurus specimen and gave priority to excavating Quarry 12, which held a more promising Morosaurus 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. 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. More Brachiosaurus fossils are reported on Riggs Hill, but other fossil finds on the hill have been vandalized. 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. In 1903, he named the type species Brachiosaurus altithorax. Latin thorax 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". No mount of Brachiosaurus 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 related Brachiosaurus brancai (now Giraffatitan) in Museum 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 of United Airlines' Terminal One in O'Hare International Airport to make room for the museum's newly acquired Tyrannosaurus skeleton, "Sue". 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. Another outdoor cast was sent to Disney'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. Assigned material Further discoveries of Brachiosaurus 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, Oklahoma, Utah, Most of the specimens collected by Felch were sent to the National Museum of Natural History in 1899 after Marsh's death, including the skull, which was then cataloged as USNM 5730. McIntosh later tentatively recognized the Felch Quarry skull as belonging to Brachiosaurus, and brought it to the attention of the American paleontologists Kenneth Carpenter and Virginia Tidwell, while urging them to describe it. They brought the skull to the Denver 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 of Giraffatitan and Camarasaurus acting as templates for the missing bones. In 2019, American paleontologists Michael D. D'Emic and Matthew T. Carrano re-examined the Felch Quarry skull after having it further prepared and CT-scanned (while consulting historical illustrations that showed earlier states of the bones), and concluded that a quadrate 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 to Camarasaurus sp. (other teeth assignable to that genus are known from the quarry). They also found it most parsimonious to assign the skull to B. 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 a phylogenetic analysis did not change the position of B. altithorax). BYU 9462 has been seen as a possible Brachiosaurus bone; it was originally assigned to Ultrasauros (now a junior synonym of Supersaurus), Museum of Ancient Life A shoulder blade with coracoid 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 exceptionally large taxa, including the new genera Supersaurus and Ultrasaurus, the latter renamed Ultrasauros shortly thereafter because another sauropod had already received the name. Later study showed that the "ultrasaur" material mostly belonged to Supersaurus, though the shoulder blade did not. Because the holotype of Ultrasauros, a dorsal vertebra, was one of the specimens that was actually from Supersaurus, the name Ultrasauros is a synonym of Supersaurus. The shoulder blade, specimen BYU 9462 (previously BYU 5001), was in 1996 assigned to a Brachiosaurus sp. (of uncertain species) by Brian Curtice and colleagues; in 2009 Michael P. Taylor concluded that it could not be referred to B. altithorax. According to Taylor in 2009, it is not clearly referable to Brachiosaurus despite its large size of . 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 to B. altithorax, as far as it is overlapping with its type specimen. Jensen further mentioned a specimen discovered near Jensen, Utah, that includes a rib in length, an anterior cervical vertebra, part of a scapula, and a coracoid, although he did not provide a description. In 2018, the largest sauropod foot ever found was reported from the Black Hills of Weston County, Wyoming. The femur is not preserved but comparisons suggest that it was about two percent longer than that of the B. altithorax holotype. Though possibly belonging to Brachiosaurus, the authors cautiously classified it as an indeterminate brachiosaurid. 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" to B. 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" with Sonorasaurus and Giraffatitan suggested brachiosaurid affinities, but this, the authors stated, would be confirmed only through further study. and 1961, Janensch compared the species in more detail, listing thirteen shared characters between Brachiosaurus brancai (which he now considered to include B. fraasi) and B. altithorax. Janensch based his description of B. brancai on "Skelett S" (skeleton S) from Tendaguru, but later realized that it comprised two partial individuals: SI and SII. He at first did not designate them as a syntype series, but in 1935 made SI (presently MB.R.2180) the lectotype. Taylor in 2009, unaware of this action, proposed the larger and more complete SII (MB.R.2181) as the lectotype. , 1915 In 1988 Gregory S. Paul published a new reconstruction of the skeleton of B. brancai, highlighting differences in proportion between it and B. 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 in B. 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 at subgenus level, as Brachiosaurus (Brachiosaurus) altithorax and Brachiosaurus (Giraffatitan) brancai. Its referral to Brachiosaurus was doubted in the 2004 edition of The Dinosauria by Paul Upchurch, Barret, and Peter Dodson who listed it as an as yet unnamed brachiosaurid genus. De Lapparent and Zbyszewski had described a series of remains but did not designate a type 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 that Lusotitan is a brachiosaurid, Based on these, Albert-Félix de Lapparent described and named the species Brachiosaurus nougaredi in 1960. He indicated the discovery locality as being in the Late Jurassic-age Taouratine Series. He assigned the rocks to this age in part because of the presumed presence of Brachiosaurus. B. nougaredi was in 2004 considered to represent a distinct, unnamed brachiosaurid genus, The metacarpals were concluded to belong to some indeterminate titanosauriform. 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 the ''Muséum national d'histoire naturelle. Only four out of the five sacral vertebrae are preserved. The total original length was in 1960 estimated at , compared to with B. altithorax. This would make it larger than any other sauropod sacrum ever found, except those of Argentinosaurus and Apatosaurus''. ==Description==

Size Most estimates of Brachiosaurus altithorax size are based on the related brachiosaurid Giraffatitan (formerly known as B. brancai), which is known from much more complete material than Brachiosaurus. The two species are the largest brachiosaurids of which relatively extensive remains have been discovered. There is another element of uncertainty for the North American Brachiosaurus because the type (and most complete) specimen appears to represent a subadult, as indicated by the unfused suture between the coracoid, a bone of the shoulder girdle that forms part of the shoulder joint, and the scapula (shoulder blade). Over the years, the mass of the holotype specimen has been estimated within the range of . but these estimates were questioned due to a very large error range and lack of precision. The length of Brachiosaurus has been estimated at 20–22 meters (66–72ft) and , While the limb bones of the most complete Giraffatitan skeleton (MB.R.2181) were very similar in size to those of the Brachiosaurus type specimen, the former was somewhat lighter than the Brachiosaurus specimen given its proportional differences. In studies including estimates for both genera, Giraffatitan was estimated at , , , Large air sacs connected to the lung system were present in the neck and trunk, invading the vertebrae and ribs by bone resorption, greatly reducing the overall density of the body. The neck is not preserved in the holotype specimen, but was very long even by sauropod standards in the closely related Giraffatitan, consisting of thirteen elongated cervical (neck) vertebrae. The neck was held in a slight S-curve, with the lower and upper sections bent and a straight middle section. Brachiosaurus and Giraffatitan probably had a small shoulder hump between the third and fifth dorsal (back) vertebra, where the sideward- and upward-directed vertebral processes were longer, providing additional surface for neck muscle attachment. The ribcage was deep compared to other sauropods. Though the humerus (upper arm bone) and femur (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 and metacarpus of other brachiosaurids. In contrast, most other sauropods had a shorter forelimb than hindlimb; the forelimb is especially short in contemporaneous diplodocoids. Brachiosaurus differed in its body proportions from the closely related Giraffatitan. The trunk was about 25 to 30 percent longer, resulting in a dorsal vertebral column longer than the humerus. Only a single complete caudal (tail) vertebra has been discovered, but its great height suggests that the tail was larger than in Giraffatitan. This vertebra had a much greater area for ligament attachment due to a broadened neural spine, indicating that the tail was also longer than in Giraffatitan, possibly by 20 to 25 percent. 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 to Giraffatitan, 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 of Giraffatitan were broader than tall. The foramina (small openings) on the sides of the centra, which allowed for the intrusion of air sacs, were larger than in Giraffatitan. The diapophyses (large projections extending sideways from the neural arch of the vertebrae) were horizontal, while those of Giraffatitan were inclined upward. At their ends, these projections articulated with the ribs; the articular surface was not distinctly triangular as in Giraffatitan. In side view, the upward-projecting neural spines stood vertically and were twice as wide at the base than at the top; those of Giraffatitan tilted backward and did not broaden at their base. When seen in front or back view, the neural spines widened toward their tops. ==Classification==

Riggs, in his preliminary 1903 description of the not yet fully prepared holotype specimen, considered Brachiosaurus to be an obvious member of the Sauropoda. To determine the validity of the genus, he compared it to the previously named genera Camarasaurus, Apatosaurus, Atlantosaurus, and Amphicoelias, 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 of Brachiosaurus itself. When describing Brachiosaurus brancai and B. fraasi in 1914, Janensch observed that the unique elongation of the humerus was shared by all three Brachiosaurus species as well as the British Pelorosaurus. He also noted this feature in Cetiosaurus, where it was not as strongly pronounced as in Brachiosaurus and Pelorosaurus. These assignments were often based on broad similarities rather than unambiguous synapomorphies, shared new traits, and most of these genera are currently regarded as dubious. In 1969, in a study by R.F. Kingham, B. altithorax, B. brancai and B. atalaiensis, along with many species now assigned to other genera, were placed in the genus Astrodon, creating an Astrodon altithorax. Kingham's views of brachiosaurid taxonomy have not been accepted by many other authors. In 1997, he published an analysis in which species traditionally considered brachiosaurids were subsequent offshoots of the stem of a larger grouping, the Titanosauriformes, and not a separate branch of their own. This study also pointed out that B. altithorax and B. brancai did not have any synapomorphies, so that there was no evidence to assume they were particularly closely related. 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. In 2009, Taylor noted multiple anatomical differences between the two Brachiosaurus species, and consequently moved B. brancai into its own genus, Giraffatitan. In contrast to earlier studies, Taylor treated both genera as distinct units in a cladistic analysis, finding them to be sister groups. Another 2010 analysis focusing on possible Asian brachiosaurid material found a clade including Abydosaurus, Brachiosaurus, Cedarosaurus, Giraffatitan, and Paluxysaurus, but not Qiaowanlong, the putative Asian brachiosaurid. Several subsequent analyses have found Brachiosaurus and Giraffatitan not to be sister groups, but instead located at different positions on the evolutionary tree. A 2012 study by D'Emic placed Giraffatitan in a more basal position, in an earlier branch, than Brachiosaurus, }} 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. ==Paleobiology==

and eyespots Habits It was believed throughout the nineteenth and early twentieth centuries that sauropods like Brachiosaurus were too massive to support their own weight on dry land, and instead lived partly submerged in water. Riggs, affirming observations by John Bell Hatcher, was the first to defend in length that most sauropods were fully terrestrial animals in his 1904 account on Brachiosaurus, 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 the zygapophyses were very reduced while the hyposphene-hypantrum complex was extremely developed, resulting in a stiff torso incapable of bending sideways. The body was fit for only quadrupedal movement on land. In 1990 the paleontologist Stephen Czerkas stated that Brachiosaurus could have entered water occasionally to cool off (thermoregulate). Neck posture Ongoing debate revolves around the neck posture of brachiosaurids, with estimates ranging from near-vertical to horizontal orientations. Reflecting this research, various newspapers ran stories criticizing the Field Museum Brachiosaurus 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). Some recent studies also advocated a more upward directed neck. Christian and Dzemski (2007) estimated that the middle part of the neck in Giraffatitan was inclined by sixty to seventy degrees; a horizontal posture could be maintained only for short periods of time. The distance between head and heart would have been reduced by the S-curvature of the neck by more than in comparison to a totally vertical posture. The neck may also have been lowered during locomotion by twenty degrees. Feeding and diet (formerly housed in the Field Museum) 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 less inclined neck, its head height may still have been over above the ground. A bipedal stance might have been adopted by Brachiosaurus in exceptional situations, like male dominance fights. The downward mobility of the neck of Brachiosaurus 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 that Brachiosaurus 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. Nostril function The bony nasal openings of neosauropods like Brachiosaurus 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 paleontologist Lawrence M. Witmer rejected this reconstruction in 2001, pointing out that all living vertebrate 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. 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 that Brachiosaurus and other sauropods with large noses could have had ornamental nasal crests. It has been proposed that sauropods, including Brachiosaurus, may have had proboscises (trunks) based on the position of the bony narial orifice, to increase their upward reach. Fabien Knoll and colleagues disputed this for Diplodocus and Camarasaurus in 2006, finding that the opening for the facial 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 for Giraffatitan was larger, and could therefore not discard the possibility of a proboscis in this genus. Metabolism Like other sauropods, Brachiosaurus was probably homeothermic (maintaining a stable internal temperature) and endothermic (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 high basic metabolic rate of its cells. Russel (1989) used Brachiosaurus as 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. 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. Another plausible ontogenetic change is the increased pneumatization of the vertebrae. During growth, the diverticula 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. ==Paleoecology==

remains from the Morrison Formation (gray); 5(the red dot) is the B. altithorax type locality Brachiosaurus is known only from the Morrison Formation of western North America (following the reassignment of the African species), from the Kimmeridgian and Tithonian ages (155.6 to 145.5 million years ago). and flat floodplains. Among these were Apatosaurus, Barosaurus, Camarasaurus, Diplodocus, Haplocanthosaurus, and Supersaurus. 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 to Barosaurus (13) and Haplocanthosaurus (12), but far fewer than Apatosaurus (112), Camarasaurus (179), and Diplodocus (98). Allosaurus accounted for 70 to 75 percent of theropod specimens and was at the top trophic level of the Morrison food web. Ceratosaurus might have specialized in attacking large sauropods, including smaller individuals of Brachiosaurus. Other vertebrates that shared this paleoenvironment included ray-finned fish, frogs, salamanders, turtles like Dorsetochelys, sphenodonts, lizards, terrestrial and aquatic crocodylomorphs such as Hoplosuchus, and several species of pterosaur like Harpactognathus and Mesadactylus. Shells of bivalves and aquatic snails are also common. The flora of the period has been revealed by fossils of green algae, fungi, mosses, horsetails, cycads, ginkgoes, and several families of conifers. Vegetation varied from river-lining forests in otherwise treeless settings (gallery forests) with tree ferns, and ferns, to fern savannas with occasional trees such as the Araucaria-like conifer Brachyphyllum. ==Cultural significance==

humerus, 1959 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. 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 local Grand Junction News announcing the find of one of the largest dinosaurs ever. On August 14, The New York Times brought the story. At the time sauropod dinosaurs appealed to the public because of their great size, often exaggerated by sensationalist newspapers. Riggs in his publications played into this by emphasizing the enormous magnitude of Brachiosaurus. Replica skeletons of Brachiosaurus can be seen in Chicago, Illinois, one outside the Field Museum and another inside the O'Hare International Airport. Brachiosaurus has been called one of the most iconic dinosaurs, but most popular depictions are based on the African species B. brancai which has since been moved to its own genus, Giraffatitan. Brachiosaurus was featured in the 1993 movie Jurassic Park, as the first computer generated dinosaur shown. The movements of the movie's Brachiosaurus were based on the gait of a giraffe combined with the mass of an elephant. A scene later in the movie used an animatronic head and neck, for when a Brachiosaurus interacts with human characters. The digital model of Brachiosaurus used in Jurassic Park later became the starting point for the ronto models in the 1997 special edition of the film Star Wars Episode IV: A New Hope. ==References==