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Giraffatitan

Giraffatitan (name meaning "titanic giraffe") is a genus of sauropod dinosaur that lived during the late Jurassic Period (KimmeridgianTithonian stages) in what is now Lindi Region, Tanzania. Only one species is known, G. brancai, named in honor of German paleontologist Wilhelm von Branca, who was a driving force behind the expedition that discovered it in the Tendaguru Formation. Giraffatitan brancai was originally described by German paleontologist Werner Janensch as a species of the North American sauropod Brachiosaurus from the Morrison Formation, as Brachiosaurus brancai. Recent research shows that the differences between the type species of Brachiosaurus and the Tendaguru material are so large that the African material should be placed in a separate genus.

Giraffatitan
Temporal range: Late Jurassic (Tithonian), 150–145 Ma
Mounted skeleton, Berlin's Natural History Museum
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Sauropodomorpha
Clade: Sauropoda
Clade: Macronaria
Family: Brachiosauridae
Genus: Giraffatitan
Paul, 1988
Type species
Giraffatitan brancai
(Janensch, 1914[1])
Synonyms
Genus synonymy[2]
  • "Abdallahsaurus"
    (nomen nudum)
  • "Blancocerosaurus"
    (nomen nudum)
  • "Ligomasaurus"
    (nomen nudum)
  • "Mtapaisaurus"
    (nomen nudum)
  • "Salimosaurus"
    (nomen nudum)
  • "Wangonisaurus"
    (nomen nudum)
Species synonymy

Giraffatitan was for many decades known as the largest dinosaur but recent discoveries of several larger dinosaurs prove otherwise; giant titanosaurians appear to have surpassed Giraffatitan in terms of sheer mass. Also, the sauropod dinosaur Sauroposeidon is estimated to be taller and possibly heavier than Giraffatitan. Most size estimates for Giraffatitan are based on the specimen HMN SII, a subadult individual, but there is evidence supporting that these animals could grow larger; specimen HMN XV2, represented by a fibula 13% larger than the corresponding material on HMN SII, would have measured around 23–26 metres (75–85 ft) long and weighed about 40–48 metric tons (44–53 short tons). Typically for a sauropod are the proportionally long neck, small skull, and large overall size of Giraffatitan. Also, the limbs were pillar-like, and the hands columnar. Unusually, in contrast, are the long forelimbs, which result in a steeply inclined trunk, making the overall body shape reminiscent of a modern giraffe. Also, while the tail is a typical long dinosaur tail, it is relatively short for a sauropod. These features show that Giraffatitan belongs in the family Brachiosauridae.

History of discovery edit

 
A hindlimb during excavation in the Tendaguru in Lindi Region

In 1906, mining engineer Bernhard Wilhelm Sattler, while travelling, noticed an enormous bone jutting out of the ground at the Tendaguru (the "steep hill") near Lindi, in what was then German East Africa, today Lindi Region, Tanzania. In early 1907, his superior Wilhelm Arning in Hannover received a report on the find. Arning again informed the Kommission für die landeskundliche Erforschung der Schutzgebiete, a commission in Berlin overviewing the geographical investigation of German protectorates.[3] The German secretary of state of colonies, Berhard Dernburg, at the time visited German East Africa accompanied by the industrialist Heinrich Otto. Otto had invited the paleontologist Professor Eberhard Fraas to join him as a scientific advisor.[4] In the summer of 1907, Fraas, already for some months travelling the colony, received a letter from Dr Hans Meyer in Leipzig urging him to investigate Sattler's discovery. On 30 August, Fraas arrived by steamer at the coastal town of Lindi.[5] A five-day march brought him to the Tendaguru, where he could confirm that the bones were authentic and dinosaurian.[6] Soon Sattler joined him with a team of native miners who uncovered two large sauropod skeletons which were transported to Germany.[7] Ultimately, these would become the holotypes of the genera Tornieria and Janenschia.

Fraas had observed that the Tendaguru layers were exceptionally rich in fossils. After his return to Germany he tried to raise enough money for a major expedition. He managed to attract the interest of Professor Wilhelm von Branca, the head of the Geologisch-Paläontologische Institut und Museum der Königliche Friedrich-Wilhelm Universität zu Berlin.[8] Von Branca considered it a matter of German national pride that such a project would succeed.[9] He involved the well-connected pathologist David von Hansemann.[10] Von Hansemann founded a Tendaguru Committee headed by Johann Albrecht, the duke of Mecklenburg. Soon it became fashionable to join this committee which counted a large number of prominent German industrialists and scientists among its members. Many of their rich friends donated considerable sums.[11] To lead the expedition, von Branca sent out one of his curators, Werner Janensch,[12] and one of his assistants, Edwin Hennig.[13] Both men arrived in Dar es Salaam on 2 April 1909.[14]

 
Porters carrying a large bone at the Tendaguru

The expedition initially employed about 160 native porters, as beasts of burden could not be used because of the danger posed by the tse tse fly. [15] During four field seasons, of 1909, 1910, 1911 and 1912, about a hundred paleontological quarries were opened. Large amounts of fossil material were shipped to Germany. Soon it became evident that apart from Tornieria and Janenschia, other sauropods were present in the layers. One was the medium-sized Dicraeosaurus, a relatively common find. More rare was a gigantic form that far surpassed the others in magnitude and that is today known as Giraffatitan. The first quarry with Giraffatitan material was "Site D", located about one kilometre northeast of the Tendaguru Hill and opened on 21 June 1909. It contained a relatively complete skeleton of a medium-sized individual, lacking the hands, the neck, the back vertebrae and the skull. It included an articulated series of twenty-nine tail vertebrae. The other bones were found in close association on a surface of twenty-two square metres. "Site IX", located 1.4 kilometres northeast of the Tendaguru Hill, was opened on 17 August 1909. Among an assemblage of 150 disarticulated dinosaur bones, also two Giraffatitan thighbones were present. The next Giraffatitan quarry was "Site N", at nine hundred metres east of the Tendaguru Hill, excavated in September 1909. It held a single disarticulated skeleton containing a back vertebra, a tail vertebra, ribs, a scapula. a possible scapula, a humerus, two ischia and a number of unidentifiable bones.[16]

The most important source of Giraffatitan fossils would be "Site S" at one kilometre southwest of the hill. Excavations started on 11 October 1909 and continued well into 1912. In 1909 limb and girdle elements were dug up. During 1910, a cut bank of the Kitukituki river was gradually deepened, removing a high overburden. To prevent the quarry walls from collapsing, they were covered by a high wooden framework. That year, first several ribs were uncovered and later part of the vertebral column. In October, close to some neck vertebrae a skull and lower jaws were discovered. From 5 June 1912 onwards more neck and trunk vertebrae were found. Initially it was thought that a single skeleton was being uncovered. Only much later Janensch realised that two skeletons had been present. Skelett SI was represented by a skull, six neck vertebrae and some back vertebrae. Skelett SII was larger but despite its size still a subadult individual. It included skull bones, a series of eleven neck and eleven back vertebrae, ribs, the left scapula, both coracoids, both forelimbs, the pubic bones and the right hindlimb. The sacrum and the tail had been lost to relatively recent erosion. The animal was found in an upright position with vertical limbs, which has been explained by its becoming mired in mud.[16]

 
Native foreman Boheti bin Amrani preparing a large rib

In early October 1909, "Site ab" was excavated, 1.2 kilometres northeast of the hill. Among disarticulated remains of many sauropods, also two Giraffatitan thighbones were collected. A gigantic possible humerus was too damaged to be salvaged. "Site cc", 2.9 kilometres northeast of the hill, contained a disarticulated Giraffatitan skeleton including neck vertebrae, a trunk vertebra, ribs, a scapula and a humerus. In 1910, another Giraffatitan quarry was opened, "Site Y" at 3.1 kilometres north of the Tendaguru Hill. It contained the skeleton of a medium-sized individual including a braincase, a series of eight neck vertebrae, a trunk vertebra, ribs, both scapulae, a coracoid, a left humerus and a left fibula.[16]

A partial skeleton called "Skelett Y" (skeleton Y) was discovered in quarry "Y". On the basis of its scapula (shoulder blade) and humerus (upper arm bone) Brachiosaurus fraasi was erected, which later turned out to be a junior synonym of B. brancai. From quarry "D" came 23 vertebrae articulated with a sacrum (a hip bone), from quarry "Aa" another 18 vertebrae with a sacrum, from quarry "no" 50 articulated tail vertebrae, from GI 16 tail vertebrae and a number of limb bones, and much more material from other places, including many limb elements.

The quarries listed above represent only the most important sites where bones were found. In dozens of other Tendaguru locations, finds were made of large single sauropod bones that were referred to the taxon in Janensch's publications but of which no field notes survive so that the precise circumstances of the discoveries are unknown. Partly this reflects a lack of systematic documentation by the expedition. Many documents were destroyed by an allied bombardment in 1943. Part of the fossils were also lost. Nevertheless, most of the skeleton is known.[17]

 
Mounted skeleton of Giraffatitan before it was remounted

Giraffatitan brancai was first named and described by German paleontologist Werner Janensch in 1914 as Brachiosaurus brancai, based on several specimens recovered between 1909 and 1912 from the Tendaguru formation.[1] It is known from five partial skeletons, including three skulls and numerous fragmentary remains including skull material, some limb bones, vertebrae and teeth. It lived from 145 to 150 million years ago, during the Kimmeridgian to Tithonian ages of the Late Jurassic period. The lectotype material of Giraffatitan brancai is skeleton "Skelett SII", a partial skeleton from Tendaguru. It consists of skull fragments including dentaries, eleven cervical vertebrae, cervical ribs, seven dorsal vertebrae, nearly complete set of dorsal ribs, distal caudal vertebrae, chevrons, left scapula, both coracoids and sternal plates, right forelimb (except the humerus, wich is probably from a referred specimen from quarry "Ni") and manus, left humerus, ulna and radius, both pubes, partial left femur, right tibia and fibula.[17] "Skelett SI" is a paralectotype, and consists of a skull and six cervical (neck) vertebrae.[17]

Originally, the Tedaguru "Skelett S" was thought by Janensch to stem from one animal, and to be very similar to the North American genus Brachiosaurus. Therefore, Janensch described it as Brachiosaurus brancai, choosing the species name in honor of Wilhelm von Branca, then director of the Museum für Naturkunde and a driving force behind the Tendaguru expedition.[1] Later, Janensch realized that the remains pertained to two separate animals, "Skelett SI" (skeleton S1) and "Skelett SII" (skeleton S2) both of which belonged to the same species.[citation needed] Janensch did not designate either of the two as lectotype, and Taylor in 2009 selected the more complete "Skellet S II" as the lectotype.[17]

A famous specimen of Giraffatitan brancai mounted in the Berlin's Natural History Museum is one of the largest, and in fact the tallest, mounted skeletons in the world, as certified by the Guinness Book of Records. Beginning in 1909, Werner Janensch found many additional G. brancai specimens in Tanzania, Africa, including some nearly complete skeletons, and used them to create the composite mounted skeleton seen today.

Separation from Brachiosaurus edit

 
Diagram incorporating bones of both Brachiosaurus and Giraffatitan, by William Diller Matthew, 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.[18] Giraffatitan was raised to full genus level by George Olshevsky in 1991, while referring to the vertebral variation.[19] Between 1991 and 2009, the name Giraffatitan was almost completely disregarded by other researchers.[17]

A detailed 2009 study by Taylor of all material, including the limb and girdle bones, found that there are significant divergences between B. 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 between Diplodocus and Barosaurus, and therefore argued that the African material should indeed be placed in its own genus (Giraffatitan) as Giraffatitan brancai.[17] An important contrast between the two genera is their overall body shape, with Brachiosaurus having a 23 percent longer dorsal vertebral series and a 20 to 25 percent longer and also taller tail.[17] The split was rejected by Daniel Chure in 2010,[20] but from 2012 onward most studies recognized the name Giraffatitan.[21]

Description edit

Size edit

 
Fossil bones in the Giraffatitan Berlin composite skeletal specimen

Between 1914 and the 1990s, Giraffatitan was claimed to be the largest dinosaur known, (ignoring the possibly larger but lost Maraapunisaurus) and thus the largest land animal in history. In the later part of the twentieth century, several giant titanosaurians found appear to surpass Giraffatitan in terms of sheer mass. However, Giraffatitan and Brachiosaurus are still the largest sauropods known from relatively complete material.[17]

All size estimates for Giraffatitan are based on the skeleton mounted in Berlin, which is partly constructed from authentic bones. These were largely taken from specimen HMN SII,[17] a subadult individual between 21.8–22.46 metres (71.5–73.7 ft) in length and about twelve meters (forty feet) tall.[22][23] The often mentioned length of 22.46 metres is by Werner Janensch, the German scientist who described Giraffatitan, and was the result of a simple adding error: the correct number should have been 22.16 metres. Mass estimates are more problematic and historically have strongly varied from as little as 15 tonnes (17 short tons) to as much as 78 tonnes (86 short tons). These extreme estimates are now considered unlikely due to flawed methodologies. There are also a large number of such estimations as the skeleton proved to be an irresistible subject for researchers wanting to test their new measuring methods. The first calculations were again made by Janensch. In 1935, he gave a volume of 32 m³ for specimen SII and of 25 m³ for specimen SI, a smaller individual. It is not known how he arrived at these numbers. In 1950, he mentioned a weight of forty tonnes for the larger skeleton.[23] In 1962, Edwin Harris Colbert measured a volume of 86.953 m³. Presuming a density of 0.9, this resulted in a weight of 78,258 kilogrammes.[24] Colbert had inserted a museum model, sold to the public, into sand and observed the volume displaced by it. Gregory S. Paul in 1988 assumed that the, in his opinion, unrealistically high number had been caused by the fact that such models used to be very bloated compared to the real build of the animal.[18] In 1980, Dale Alan Russell et al published a much lower weight of 14.8 tonnes by extrapolating from the diameter of the humerus and the thighbone.[25] In 1985, the same researcher arrived at 29 tonnes by extrapolating from the circumference of these bones.[26] In 1985, Robert McNeill Alexander found a value of 46.6 tonnes inserting a toy model of the British Museum of Natural History into water.[27]

More recent estimates based on models reconstructed from bone volume measurements, which take into account the extensive, weight-reducing airsac systems present in sauropods, and estimated muscle mass, are in the range of 23.3–39.5 tonnes (25.7–43.5 short tons).[17][22] In 1988, G.S. Paul measured a volume of 36.585 m³ by inserting a specially constructed model into water. He estimated a weight of 31.5 tonnes, assuming a low density.[18] In 1994/1995 a weight of 40 tonnes extrapolating from limb bone circumference.[28] In 1995 a laser scan of the skeleton was used to build a virtual model from simple geometrical shapes, finding a volume of 74.42 m³ and concluding to a weight of 63 tonnes.[29] In 2008, Gunga revised the volume, using more complex shapes, to 47.9 m³.[30] Donald Henderson in 2004 employed a computer model that calculated a volume of 32.398 m³ and a weight of 25,789 kilogrammes.[31] Newer methods use bone wall thickness.[32]

However, HMN SII is not the largest specimen known (an assertion supported by its subadult status) but HMN XV2, represented by a fibula 13% larger than the corresponding material on HMN SII,[17] which might have attained 26 metres (85 ft) in length.[33] Gregory S. Paul initially estimated the size of this specimen at 25 metres (82 ft) in total length, 16 metres (52 ft) in total height and 45 metric tons (50 short tons) in body mass,[18] but later moderated at 23 metres (75 ft) in total length and 40 metric tons (44 short tons) in body mass.[34] In 2020, Molina-Perez and Larramnedi estimated the size of the HMN XV2 specimen at 25 metres (82 ft) and 48 tonnes (53 short tons), with a shoulder height of 6.8 metres (22 ft).[2]

General build edit

 
Restoration

Giraffatitan was a sauropod, one of a group of four-legged, plant-eating dinosaurs with long necks and tails and relatively small brains. It had a giraffe-like build, with long forelimbs and a very long neck. The skull had a tall arch anterior to the eyes, consisting of the bony nares, a number of other openings, and "spatulate" teeth (resembling chisels). The first toe on its front foot and the first three toes on its hind feet were clawed.

Skull edit

 
Skull cast in Berlin

Traditionally, the distinctive high-crested skull was seen as a characteristic of the genus Brachiosaurus, to which Giraffatitan brancai was originally referred; however, it is possible that Brachiosaurus altithorax did not show this feature, since within the traditional Brachiosaurus material it is known only from Tanzanian specimens now assigned to Giraffatitan.

The placement of Giraffatitan nostrils has been the source of much debate with Witmer (2001) describing in Science the hypothesized position of the fleshy nostrils in Giraffatitan in as many as five possible locations. Comparing the nares of dinosaurs with those of modern animals, he found that all species have their external nostril openings in the front, and that sauropods like Giraffatitan did not have nostrils on top of their heads, but near their snouts.[35] There has also been the hypothesis of various sauropods, such as Giraffatitan, possessing a trunk. The fact that there were no narrow-snouted sauropods (Giraffatitan included) tends to discredit such a hypothesis. Stronger evidence for the absence of a trunk is found in the teeth wear of Giraffatitan, which shows the kind of wear that would result from biting and tearing off of plant matter rather than purely grinding, which would be the result of having already ripped the leaves and branches off with a trunk.[36]

Classification edit

 
Reconstructed skeleton of the closely related Brachiosaurus

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.[1] Janensch concluded that the four genera must have been closely related to each other, and in 1929 assigned them to a subfamily Brachiosaurinae within the family Bothrosauropodidae.[37]

During the twentieth century, several sauropods were assigned to Brachiosauridae, including Astrodon, Bothriospondylus, Pelorosaurus, Pleurocoelus, and Ultrasauros.[38] 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.[39][40] 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.[41] Kingham's views of brachiosaurid taxonomy have not been accepted by many other authors.[42] Since the 1990s, computer-based cladistic analyses allow for postulating detailed hypotheses on the relationships between species, by calculating those trees 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.[43] 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.[44]

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.[45] The exact status of each potential brachiosaurid varies from study to study. For example, a 2010 study by Chure and colleagues recognized Abydosaurus as a brachiosaurid together with Brachiosaurus, which in this study included B. brancai.[20] 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.[45] 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,[42] while a 2013 study by Philip Mannion and colleagues had it the other way around.[21]

A 2012 study on titanosauriform sauropods by Michael D'Emic placed Giraffatitan as sister to a clade containing Brachiosaurus and a tritomy of Abydosaurus, Cedarosaurus, and Venenosaurus as shown in the cladogram below:[46]

 
Authentic Giraffatitan skull (behind), compared to that of the small Europasaurus

In their 2024 description of Gandititan, Han et al. analyzed the phylogenetic relations of Macronaria, focusing on titanosauriform taxa. They recovered Giraffatitan as the sister taxon to Sonorasaurus, similar to some previous analyses,[47] in a clade also containing Brachiosaurus. The results of their phylogenetic analyses are shown in the cladogram below:[48]

Paleobiology edit

 
The feeding range of Giraffatitan (left) and Sauroposeidon

The nostrils of Giraffatitan, like the huge corresponding nasal openings in its skull, were long thought to be located on the top of the head. In past decades, scientists theorized that the animal used its nostrils like a snorkel, spending most of its time submerged in water in order to support its great mass. The current consensus view, however, is that Giraffatitan was a fully terrestrial animal. Studies have demonstrated that water pressure would have prevented the animal from breathing effectively while submerged and that its feet were too narrow for efficient aquatic use. Furthermore, new studies by Lawrence Witmer (2001) show that, while the nasal openings in the skull were placed high above the eyes, the nostrils would still have been close to the tip of the snout (a study which also lends support to the idea that the tall "crests" of brachiosaurs supported some sort of fleshy resonating chamber).

Brain edit

Giraffatitan's brain measured about 300 cubic centimetres, which, like those of other sauropods, was small compared to its massive body size. A 2009 study calculated its Encephalization Quotient (a rough estimate of possible intelligence) at a low 0.62 or 0.79, depending on the size estimate used. Like other sauropods, Giraffatitan has a sacral enlargement above the hip which some older sources misleadingly referred to as a "second brain".[49] However, glycogen bodies are a more likely explanation.[50]

Nostril function edit

 
The fleshy external nostril would have been placed at the front of the nasal fossa, the depression seen in front of the bony nostril

The bony nasal openings of neosauropods like Giraffatitan 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.[51]

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.[52]

It has been proposed that sauropods, including Giraffatitan, 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.[53]

Metabolism edit

If Giraffatitan was endothermic (warm-blooded), it would have taken an estimated ten years to reach full size; if it were instead poikilothermic (cold-blooded), then it would have required over 100 years to reach full size.[54] As a warm-blooded animal, the daily energy demands of Giraffatitan would have been enormous; it would probably have needed to eat more than ~182 kg (400 lb) of food per day. If Giraffatitan was fully cold-blooded or was a passive bulk endotherm, it would have needed far less food to meet its daily energy needs. Some scientists have proposed that large dinosaurs like Giraffatitan were gigantotherms.[55] Internal organs of these giant sauropods were probably enormous.[56]

Paleoenvironment edit

 
Restoration of two individuals

Giraffatitan lived in what is now Tanzania in the Late Jurassic Tendaguru Formation.[57] Since 2012, the boundary between the Kimmeridgian and Tithonian is dated at 152.1 million years ago.[58]

The Tendaguru ecosystem primarily consisted of three types of environment: shallow, lagoon-like marine environments, tidal flats and low coastal environments; and vegetated inland environments. The marine environment existed above the fair weather wave base and behind siliciclastic and ooid barriers. It appeared to have had little change in salinity levels and experienced tides and storms. The coastal environments consisted of brackish coastal lakes, ponds and pools. These environments had little vegetation and were probably visited by herbivorous dinosaurs mostly during droughts. The well vegetated inlands were dominated by conifers. Overall, the Late Jurassic Tendaguru climate was subtropical to tropical with seasonal rains and pronounced dry periods. During the Early Cretaceous, the Tendaguru became more humid.[59] The Tendaguru Beds are similar to the Morrison Formation of North America except in its marine interbeds.[60]

Giraffatitan would have coexisted with fellow sauropods like Dicraeosaurus hansemanni and D. sattleri, Janenschia africana, Tendaguria tanzaniensis and Tornieria africanus; ornithischians like Dysalotosaurus lettowvorbecki and Kentrosaurus aethiopicus; the theropods "Allosaurus" tendagurensis, "Ceratosaurus" roechlingi, "Ceratosaurus" ingens, Elaphrosaurus bambergi, Veterupristisaurus milneri and Ostafrikasaurus crassiserratus; and the pterosaur Tendaguripterus recki.[61][62][63][64] Other organisms that inhabited the Tendaguru included corals, echinoderms, cephalopods, bivalves, gastropods, decapods, sharks, neopterygian fish, crocodilians and small mammals like Brancatherulum tendagurensis.[65]

See also edit

References edit

  1. ^ a b c d e Janensch, W (1914). "Übersicht über der Wirbeltierfauna der Tendaguru-Schichten nebst einer kurzen Charakterisierung der neu aufgeführten Arten von Sauropoden". Archiv für Biontologie. 3 (1): 81–110.
  2. ^ a b Molina-Perez & Larramendi (2020). Dinosaur Facts and Figures: The Sauropods and Other Sauropodomorphs. New Jersey: Princeton University Press. p. 259.
  3. ^ Maier (2003), p 1
  4. ^ Maier (2003), p 3
  5. ^ Maier (2003), p 8
  6. ^ Maier (2003), p 10
  7. ^ Maier (2003), p 11-12
  8. ^ Maier (2003), p 15
  9. ^ Maier (2003), p 16
  10. ^ Maier (2003), p 17
  11. ^ Maier (2003), p 18
  12. ^ Maier (2003), p 19
  13. ^ Maier (2003), p 22
  14. ^ Maier (2003), p 23
  15. ^ Tamborini, Marco & Mareike Vennen (2017) Disruptions and changing habits: The case of the Tendaguru expedition, Museum History Journal, 10:2, 183-199.
  16. ^ a b c Heinrich, Wolf-Dieter (1999). "The Taphonomy of Dinosaurs from the Upper Jurassic of Tendaguru (Tanzania) Based on Field Sketches of the German Tendaguru Expedition (1909-1913)". Mitteilungen aus dem Museum für Naturkunde in Berlin, Geowissenschaftliche Reihe. 2: 25–61. doi:10.5194/fr-2-25-1999.
  17. ^ a b c d e f g h i j k Taylor, M.P. (2009). "A Re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropod) and its generic separation from Giraffatitan brancai (Janensch 1914)" (PDF). Journal of Vertebrate Paleontology. 29 (3): 787–806. doi:10.1671/039.029.0309. S2CID 15220647.
  18. ^ a b c d Paul, G.S. (1988). "The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world's largest dinosaurs" (PDF). Hunteria. 2 (3): 1–14. (PDF) from the original on 27 June 2022.
  19. ^ Olshevsky, G. (1991). (PDF). Mesozoic Meanderings. 2: 1–196. Archived from the original (PDF) on 19 August 2018. Retrieved 14 April 2018.
  20. ^ a b Chure, D.; Britt, B.; Whitlock, J. A.; Wilson, J. A. (2010). "First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition". Naturwissenschaften. 97 (4): 379–391. Bibcode:2010NW.....97..379C. doi:10.1007/s00114-010-0650-6. PMC 2841758. PMID 20179896.
  21. ^ a b Mannion, P. D.; Upchurch, Paul; Barnes, Rosie N.; Mateus, Octávio (2013). "Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms" (PDF). Zoological Journal of the Linnean Society. 168: 98–206. doi:10.1111/zoj.12029.
  22. ^ a b Mazzetta, G.V.; et al. (2004). "Giants and Bizarres: Body Size of Some Southern South American Cretaceous Dinosaurs". Historical Biology. 16 (2–4): 1–13. CiteSeerX 10.1.1.694.1650. doi:10.1080/08912960410001715132. S2CID 56028251.
  23. ^ a b Janensch, W. (1950). "The Skeleton Reconstruction of Brachiosaurus brancai" (PDF). Palaeontographica: 97–103.
  24. ^ Colbert, E (1962). "The weights of dinosaurs". American Museum Novitates (2076): 1–16.
  25. ^ Russell, D.; Béland, P.; McIntosh, J.S. (1980). "Paleoecology of the dinosaurs of Tendaguru (Tanzania)". Mémoires de la Société Géologique de France. 59: 169–175.
  26. ^ Anderson, J.F.; Hall-Martin, A.; Russell, D.A. (1985). "Long-bone circumference and weight in mammals, birds and dinosaurs". Journal of Zoology. 207 (1): 53–61. doi:10.1111/j.1469-7998.1985.tb04915.x.
  27. ^ Alexander, R. McN. (1985). "Mechanics of posture and gait of some large dinosaurs". Zoological Journal of the Linnean Society. 83 (1): 1–25. doi:10.1111/j.1096-3642.1985.tb00871.x.
  28. ^ Peczkis, J. (1995). "Implications of body-mass estimates for dinosaurs". Journal of Vertebrate Paleontology. 14 (4): 520–533. doi:10.1080/02724634.1995.10011575. JSTOR 4523591.
  29. ^ Gunga, H.-C.; Kirsch, K.A.; Baartz, F.; Röcker, L.; Heinrich, W.-D.; Lisowski, W.; Wiedemann, A.; Albertz, J. (1995). "New data on the dimensions of Brachiosaurus brancai and their physiological implications". Naturwissenschaften. 82 (4): 190–192. Bibcode:1995NW.....82..190G. doi:10.1007/s001140050167.
  30. ^ Gunga, H.-C.; Suthau, T.; Bellmann, A.; Stoinski, S.; Friedrich, A.; Trippel, T.; Kirsch, K.; Hellwich, O. (2008). "A new body mass estimation of Brachiosaurus brancai Janensch, 1914 mounted and exhibited at the Museum of Natural History (Berlin, Germany)". Fossil Record. 11 (1): 33–38. doi:10.1002/mmng.200700011.
  31. ^ Henderson, D.M. (2004). "Tipsy Punters: Sauropod Dinosaur Pneumaticity, Buoyancy and Aquatic Habits". Proceedings: Biological Sciences. 271 (Supplement 4): 180–183. doi:10.1098/rsbl.2003.0136. JSTOR 4142711. PMC 1810024. PMID 15252977. S2CID 23454835.
  32. ^ Benson, R. B. J.; Campione, N. S. E.; Carrano, M. T.; Mannion, P. D.; Sullivan, C.; Upchurch, P.; Evans, D. C. (2014). "Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage". PLOS Biology. 12 (5): e1001853. doi:10.1371/journal.pbio.1001853. PMC 4011683. PMID 24802911. S2CID 16100066.
  33. ^ Holtz, Thomas R. Jr. (2008) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages Supplementary Information
  34. ^ Paul, Gregory S. (2016). The Princeton Field Guide to Dinosaurs. Princeton University Press. p. 227. ISBN 978-1-78684-190-2. OCLC 985402380.
  35. ^ Witmer, L.M. (2001). (PDF). Science. 293 (5531): 850–853. CiteSeerX 10.1.1.629.1744. doi:10.1126/science.1062681. PMID 11486085. S2CID 7328047. Archived from the original (PDF) on 6 September 2013.
  36. ^ Naish, D. (20 March 2009). . ScienceBlogs. Archived from the original on 13 January 2012.
  37. ^ Janensch, 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.
  38. ^ McIntosh, John; Osmólska, H. (1990). "Sauropoda". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). The Dinosauria (1 ed.). Berkeley: University of California Press. p. 376. ISBN 978-0-520-06726-4.
  39. ^ Upchurch, P.; Barrett, P.M.; Dodson, P. (2004). "Sauropoda". In Weishampel, D.B.; Dodson, P.; Osmolska, H. (eds.). The Dinosauria, Second Edition. Univ of California Press, Berkeley. pp. 259–322. ISBN 978-0-520-24209-8.
  40. ^ Wedel, Mathew J.; Cifelli, R. L.; Sanders, R. K. (2000). "Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon". Acta Palaeontologica Polonica. 45: 343–388.
  41. ^ Kingham, R.F. (1962). "Studies of the sauropod dinosaur Astrodon Leidy". Proceedings of the Washington Junior Academy of Sciences. 1: 38–44.
  42. ^ a b D'Emic, M. D. (2012). "The early evolution of titanosauriform sauropod dinosaurs". Zoological Journal of the Linnean Society. 166 (3): 624–671. doi:10.1111/j.1096-3642.2012.00853.x. hdl:2027.42/94293.
  43. ^ Leonardo Salgado, 1993, "Comments on Chubutisaurus insignis del Corro (Saurischia, Sauropoda)", Ameghiniana 30(3): 265–270
  44. ^ Salgado, L., R. A. Coria, and J. O. Calvo. 1997. "Evolution of titanosaurid sauropods. I: phylogenetic analysis based on the postcranial evidence". Ameghiniana 34: 3-32
  45. ^ a b Ksepka, D. T.; Norell, M. A. (2010). "The illusory evidence for Asian Brachiosauridae: new material of Erketu ellisoni and a phylogenetic appraisal of basal Titanosauriformes" (PDF). American Museum Novitates (3700): 1–27. doi:10.1206/3700.2. S2CID 86254470. Archived from the original (PDF) on 19 August 2018.
  46. ^ D'emic, Michael D. (2012). "The early evolution of titanosauriform sauropod dinosaurs" (PDF). Zoological Journal of the Linnean Society. 166 (3): 624–671. doi:10.1111/j.1096-3642.2012.00853.x. hdl:2027.42/94293. S2CID 54752135.
  47. ^ Philip D. Mannion; Ronan Allain; Olivier Moine (2017). "The earliest known titanosauriform sauropod dinosaur and the evolution of Brachiosauridae". PeerJ. 5: e3217. doi:10.7717/peerj.3217. PMC 5417094. PMID 28480136.
  48. ^ Han, F.; Yang, L.; Lou, F.; Sullivan, C.; Xu, X.; Qiu, W.; Liu, H.; Yu, J.; Wu, R.; Ke, Y.; Xu, M.; Hu, J.; Lu, P. (2024). "A new titanosaurian sauropod, Gandititan cavocaudatus gen. et sp. nov., from the Late Cretaceous of southern China". Journal of Systematic Palaeontology. 22 (1). 2293038. doi:10.1080/14772019.2023.2293038.
  49. ^ Knoll, F.; Schwarz-Wings, D. (2009). "Palaeoneuroanatomy of Brachiosaurus". Annales de Paléontologie. 95 (3): 165–175. doi:10.1016/j.annpal.2009.06.001.
  50. ^ Giffin, Emily B. (1990). "Gross Spinal Anatomy and Limb Use in Living and Fossil Reptiles". Paleobiology. 16 (4): 448–485. doi:10.1017/S0094837300010186. ISSN 0094-8373. JSTOR 2400969.
  51. ^ Witmer, L. M. (2001). "Nostril position in dinosaurs and other vertebrates and its significance for nasal function". Science. 293 (5531): 850–853. CiteSeerX 10.1.1.629.1744. doi:10.1126/science.1062681. PMID 11486085. S2CID 7328047.
  52. ^ Czerkas, S. J.; Czerkas, S. A. (1990). Dinosaurs: a Global View. Limpsfield: Dragons' World. pp. 134–135. ISBN 978-0-7924-5606-3.
  53. ^ Knoll, F.; Galton, P. M.; López-Antoñanzas, R. (2006). "Paleoneurological evidence against a proboscis in the sauropod dinosaur Diplodocus". Geobios. 39 (2): 215–221. Bibcode:2006Geobi..39..215K. doi:10.1016/j.geobios.2004.11.005.
  54. ^ Case, T.J. (1978). "Speculations on the Growth Rate and Reproduction of Some Dinosaurs". Paleobiology. 4 (3): 323. doi:10.1017/S0094837300006023.
  55. ^ Bailey, J.B. (1997). "Neural spine elongation in dinosaurs: Sailbacks or buffalo-backs?". Journal of Paleontology. 71 (6): 1124–1146. doi:10.1017/s0022336000036076. S2CID 130861276.
  56. ^ "Jak velké vnitřní orgány měli obří sauropodi?". 4 July 2016.
  57. ^ Bussert, Robert; Heinrich, Wolf-Dieter; Aberhan, Martin (2009). "The Tendaguru Formation (Late Jurassic to Early Cretaceous, southern Tanzania): definition, palaeoenvironments, and sequence stratigraphy". Fossil Record. 12 (2): 141–174. doi:10.1002/mmng.200900004.
  58. ^ Gradstein, F.M.; Ogg, J.G.; Schmitz, M.D. & Ogg, G.M., 2012, A Geologic Time Scale 2012, Elsevier
  59. ^ Aberhan, Martin; Bussert, Robert; Heinrich, Wolf-Dieter; Schrank, Eckhart; Schultka, Stephan; Sames, Benjamin; Kriwet, Jürgen; Kapilima, Saidi (2002). "Palaeoecology and depositional environments of the Tendaguru Beds (Late Jurassic to Early Cretaceous, Tanzania)". Fossil Record. 5 (1): 19–44. doi:10.1002/mmng.20020050103.
  60. ^ Mateus, Octávio (2006). "Late Jurassic dinosaurs from the Morrison Formation (USA), the Lourinhā and Alcobaça formations (Portugal), and the Tendaguru Beds (Tanzania): a comparison". In Foster, J.R.; Lucas, S.G. (eds.). Paleontology and Geology of the Upper Morrison Formation. New Mexico Museum of Natural History and Science (bulletin 36). pp. 223–232. ISSN 1524-4156. {{cite book}}: |journal= ignored (help)
  61. ^ Weishampel, David B; et al. (2004). "Dinosaur distribution (Late Jurassic, Africa)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. p. 552. ISBN 0-520-24209-2.
  62. ^ Barrett, P.M., Butler, R.J., Edwards, N.P., & Milner, A.R. Pterosaur distribution in time and space: an atlas. p61–107. in Flugsaurier: Pterosaur papers in honour of Peter Wellnhofer. 2008. Hone, D.W.E., and Buffetaut, E. (eds). Zitteliana B, 28. 264pp.[1]
  63. ^ Rauhut, Oliver W. M. (2011). "Theropod dinosaurs from the Late Jurassic of Tendaguru (Tanzania)". Special Papers in Palaeontology. 86: 195–239.
  64. ^ Buffetaut, Eric (2012). "An early spinosaurid dinosaur from the Late Jurassic of Tendaguru (Tanzania) and the evolution of the spinosaurid dentition". Oryctos. 10: 1–8.
  65. ^ Heinrich, Wolf-Dieter; et al. (2001). "The German‐Tanzanian Tendaguru Expedition 2000". Fossil Record. 4 (1): 223–237. doi:10.1002/mmng.20010040113.

Bibliography edit

  • Maier, Gerhard. 2003. African dinosaurs unearthed: the Tendaguru expeditions. Life of the Past Series (ed. J. Farlow). Indiana University Press, Bloomington, Indiana

giraffatitan, name, meaning, titanic, giraffe, genus, sauropod, dinosaur, that, lived, during, late, jurassic, period, kimmeridgian, tithonian, stages, what, lindi, region, tanzania, only, species, known, brancai, named, honor, german, paleontologist, wilhelm,. Giraffatitan name meaning titanic giraffe is a genus of sauropod dinosaur that lived during the late Jurassic Period Kimmeridgian Tithonian stages in what is now Lindi Region Tanzania Only one species is known G brancai named in honor of German paleontologist Wilhelm von Branca who was a driving force behind the expedition that discovered it in the Tendaguru Formation Giraffatitan brancai was originally described by German paleontologist Werner Janensch as a species of the North American sauropod Brachiosaurus from the Morrison Formation as Brachiosaurus brancai Recent research shows that the differences between the type species of Brachiosaurus and the Tendaguru material are so large that the African material should be placed in a separate genus GiraffatitanTemporal range Late Jurassic Tithonian 150 145 Ma PreꞒ Ꞓ O S D C P T J K Pg N Mounted skeleton Berlin s Natural History MuseumScientific classificationDomain EukaryotaKingdom AnimaliaPhylum ChordataClade DinosauriaClade SaurischiaClade SauropodomorphaClade SauropodaClade MacronariaFamily BrachiosauridaeGenus GiraffatitanPaul 1988Type species Giraffatitan brancai Janensch 1914 1 SynonymsGenus synonymy 2 Abdallahsaurus nomen nudum Blancocerosaurus nomen nudum Ligomasaurus nomen nudum Mtapaisaurus nomen nudum Salimosaurus nomen nudum Wangonisaurus nomen nudum Species synonymy Brachiosaurus brancaiJanensch 1914Brachiosaurus fraasiJanensch 1914 1 Giraffatitan was for many decades known as the largest dinosaur but recent discoveries of several larger dinosaurs prove otherwise giant titanosaurians appear to have surpassed Giraffatitan in terms of sheer mass Also the sauropod dinosaur Sauroposeidon is estimated to be taller and possibly heavier than Giraffatitan Most size estimates for Giraffatitan are based on the specimen HMN SII a subadult individual but there is evidence supporting that these animals could grow larger specimen HMN XV2 represented by a fibula 13 larger than the corresponding material on HMN SII would have measured around 23 26 metres 75 85 ft long and weighed about 40 48 metric tons 44 53 short tons Typically for a sauropod are the proportionally long neck small skull and large overall size of Giraffatitan Also the limbs were pillar like and the hands columnar Unusually in contrast are the long forelimbs which result in a steeply inclined trunk making the overall body shape reminiscent of a modern giraffe Also while the tail is a typical long dinosaur tail it is relatively short for a sauropod These features show that Giraffatitan belongs in the family Brachiosauridae Contents 1 History of discovery 1 1 Separation from Brachiosaurus 2 Description 2 1 Size 2 2 General build 2 3 Skull 3 Classification 4 Paleobiology 4 1 Brain 4 2 Nostril function 4 3 Metabolism 5 Paleoenvironment 6 See also 7 References 8 BibliographyHistory of discovery edit nbsp A hindlimb during excavation in the Tendaguru in Lindi RegionIn 1906 mining engineer Bernhard Wilhelm Sattler while travelling noticed an enormous bone jutting out of the ground at the Tendaguru the steep hill near Lindi in what was then German East Africa today Lindi Region Tanzania In early 1907 his superior Wilhelm Arning in Hannover received a report on the find Arning again informed the Kommission fur die landeskundliche Erforschung der Schutzgebiete a commission in Berlin overviewing the geographical investigation of German protectorates 3 The German secretary of state of colonies Berhard Dernburg at the time visited German East Africa accompanied by the industrialist Heinrich Otto Otto had invited the paleontologist Professor Eberhard Fraas to join him as a scientific advisor 4 In the summer of 1907 Fraas already for some months travelling the colony received a letter from Dr Hans Meyer in Leipzig urging him to investigate Sattler s discovery On 30 August Fraas arrived by steamer at the coastal town of Lindi 5 A five day march brought him to the Tendaguru where he could confirm that the bones were authentic and dinosaurian 6 Soon Sattler joined him with a team of native miners who uncovered two large sauropod skeletons which were transported to Germany 7 Ultimately these would become the holotypes of the genera Tornieria and Janenschia Fraas had observed that the Tendaguru layers were exceptionally rich in fossils After his return to Germany he tried to raise enough money for a major expedition He managed to attract the interest of Professor Wilhelm von Branca the head of the Geologisch Palaontologische Institut und Museum der Konigliche Friedrich Wilhelm Universitat zu Berlin 8 Von Branca considered it a matter of German national pride that such a project would succeed 9 He involved the well connected pathologist David von Hansemann 10 Von Hansemann founded a Tendaguru Committee headed by Johann Albrecht the duke of Mecklenburg Soon it became fashionable to join this committee which counted a large number of prominent German industrialists and scientists among its members Many of their rich friends donated considerable sums 11 To lead the expedition von Branca sent out one of his curators Werner Janensch 12 and one of his assistants Edwin Hennig 13 Both men arrived in Dar es Salaam on 2 April 1909 14 nbsp Porters carrying a large bone at the TendaguruThe expedition initially employed about 160 native porters as beasts of burden could not be used because of the danger posed by the tse tse fly 15 During four field seasons of 1909 1910 1911 and 1912 about a hundred paleontological quarries were opened Large amounts of fossil material were shipped to Germany Soon it became evident that apart from Tornieria and Janenschia other sauropods were present in the layers One was the medium sized Dicraeosaurus a relatively common find More rare was a gigantic form that far surpassed the others in magnitude and that is today known as Giraffatitan The first quarry with Giraffatitan material was Site D located about one kilometre northeast of the Tendaguru Hill and opened on 21 June 1909 It contained a relatively complete skeleton of a medium sized individual lacking the hands the neck the back vertebrae and the skull It included an articulated series of twenty nine tail vertebrae The other bones were found in close association on a surface of twenty two square metres Site IX located 1 4 kilometres northeast of the Tendaguru Hill was opened on 17 August 1909 Among an assemblage of 150 disarticulated dinosaur bones also two Giraffatitan thighbones were present The next Giraffatitan quarry was Site N at nine hundred metres east of the Tendaguru Hill excavated in September 1909 It held a single disarticulated skeleton containing a back vertebra a tail vertebra ribs a scapula a possible scapula a humerus two ischia and a number of unidentifiable bones 16 The most important source of Giraffatitan fossils would be Site S at one kilometre southwest of the hill Excavations started on 11 October 1909 and continued well into 1912 In 1909 limb and girdle elements were dug up During 1910 a cut bank of the Kitukituki river was gradually deepened removing a high overburden To prevent the quarry walls from collapsing they were covered by a high wooden framework That year first several ribs were uncovered and later part of the vertebral column In October close to some neck vertebrae a skull and lower jaws were discovered From 5 June 1912 onwards more neck and trunk vertebrae were found Initially it was thought that a single skeleton was being uncovered Only much later Janensch realised that two skeletons had been present Skelett SI was represented by a skull six neck vertebrae and some back vertebrae Skelett SII was larger but despite its size still a subadult individual It included skull bones a series of eleven neck and eleven back vertebrae ribs the left scapula both coracoids both forelimbs the pubic bones and the right hindlimb The sacrum and the tail had been lost to relatively recent erosion The animal was found in an upright position with vertical limbs which has been explained by its becoming mired in mud 16 nbsp Native foreman Boheti bin Amrani preparing a large ribIn early October 1909 Site ab was excavated 1 2 kilometres northeast of the hill Among disarticulated remains of many sauropods also two Giraffatitan thighbones were collected A gigantic possible humerus was too damaged to be salvaged Site cc 2 9 kilometres northeast of the hill contained a disarticulated Giraffatitan skeleton including neck vertebrae a trunk vertebra ribs a scapula and a humerus In 1910 another Giraffatitan quarry was opened Site Y at 3 1 kilometres north of the Tendaguru Hill It contained the skeleton of a medium sized individual including a braincase a series of eight neck vertebrae a trunk vertebra ribs both scapulae a coracoid a left humerus and a left fibula 16 A partial skeleton called Skelett Y skeleton Y was discovered in quarry Y On the basis of its scapula shoulder blade and humerus upper arm bone Brachiosaurus fraasi was erected which later turned out to be a junior synonym of B brancai From quarry D came 23 vertebrae articulated with a sacrum a hip bone from quarry Aa another 18 vertebrae with a sacrum from quarry no 50 articulated tail vertebrae from GI 16 tail vertebrae and a number of limb bones and much more material from other places including many limb elements The quarries listed above represent only the most important sites where bones were found In dozens of other Tendaguru locations finds were made of large single sauropod bones that were referred to the taxon in Janensch s publications but of which no field notes survive so that the precise circumstances of the discoveries are unknown Partly this reflects a lack of systematic documentation by the expedition Many documents were destroyed by an allied bombardment in 1943 Part of the fossils were also lost Nevertheless most of the skeleton is known 17 nbsp Mounted skeleton of Giraffatitan before it was remountedGiraffatitan brancai was first named and described by German paleontologist Werner Janensch in 1914 as Brachiosaurus brancai based on several specimens recovered between 1909 and 1912 from the Tendaguru formation 1 It is known from five partial skeletons including three skulls and numerous fragmentary remains including skull material some limb bones vertebrae and teeth It lived from 145 to 150 million years ago during the Kimmeridgian to Tithonian ages of the Late Jurassic period The lectotype material of Giraffatitan brancai is skeleton Skelett SII a partial skeleton from Tendaguru It consists of skull fragments including dentaries eleven cervical vertebrae cervical ribs seven dorsal vertebrae nearly complete set of dorsal ribs distal caudal vertebrae chevrons left scapula both coracoids and sternal plates right forelimb except the humerus wich is probably from a referred specimen from quarry Ni and manus left humerus ulna and radius both pubes partial left femur right tibia and fibula 17 Skelett SI is a paralectotype and consists of a skull and six cervical neck vertebrae 17 Originally the Tedaguru Skelett S was thought by Janensch to stem from one animal and to be very similar to the North American genus Brachiosaurus Therefore Janensch described it as Brachiosaurus brancai choosing the species name in honor of Wilhelm von Branca then director of the Museum fur Naturkunde and a driving force behind the Tendaguru expedition 1 Later Janensch realized that the remains pertained to two separate animals Skelett SI skeleton S1 and Skelett SII skeleton S2 both of which belonged to the same species citation needed Janensch did not designate either of the two as lectotype and Taylor in 2009 selected the more complete Skellet S II as the lectotype 17 A famous specimen of Giraffatitan brancai mounted in the Berlin s Natural History Museum is one of the largest and in fact the tallest mounted skeletons in the world as certified by the Guinness Book of Records Beginning in 1909 Werner Janensch found many additional G brancai specimens in Tanzania Africa including some nearly complete skeletons and used them to create the composite mounted skeleton seen today Separation from Brachiosaurus edit nbsp Diagram incorporating bones of both Brachiosaurus and Giraffatitan by William Diller Matthew 1915In 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 18 Giraffatitan was raised to full genus level by George Olshevsky in 1991 while referring to the vertebral variation 19 Between 1991 and 2009 the name Giraffatitan was almost completely disregarded by other researchers 17 A detailed 2009 study by Taylor of all material including the limb and girdle bones found that there are significant divergences between B 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 between Diplodocus and Barosaurus and therefore argued that the African material should indeed be placed in its own genus Giraffatitan as Giraffatitan brancai 17 An important contrast between the two genera is their overall body shape with Brachiosaurus having a 23 percent longer dorsal vertebral series and a 20 to 25 percent longer and also taller tail 17 The split was rejected by Daniel Chure in 2010 20 but from 2012 onward most studies recognized the name Giraffatitan 21 Description editSize edit nbsp Fossil bones in the Giraffatitan Berlin composite skeletal specimenBetween 1914 and the 1990s Giraffatitan was claimed to be the largest dinosaur known ignoring the possibly larger but lost Maraapunisaurus and thus the largest land animal in history In the later part of the twentieth century several giant titanosaurians found appear to surpass Giraffatitan in terms of sheer mass However Giraffatitan and Brachiosaurus are still the largest sauropods known from relatively complete material 17 All size estimates for Giraffatitan are based on the skeleton mounted in Berlin which is partly constructed from authentic bones These were largely taken from specimen HMN SII 17 a subadult individual between 21 8 22 46 metres 71 5 73 7 ft in length and about twelve meters forty feet tall 22 23 The often mentioned length of 22 46 metres is by Werner Janensch the German scientist who described Giraffatitan and was the result of a simple adding error the correct number should have been 22 16 metres Mass estimates are more problematic and historically have strongly varied from as little as 15 tonnes 17 short tons to as much as 78 tonnes 86 short tons These extreme estimates are now considered unlikely due to flawed methodologies There are also a large number of such estimations as the skeleton proved to be an irresistible subject for researchers wanting to test their new measuring methods The first calculations were again made by Janensch In 1935 he gave a volume of 32 m for specimen SII and of 25 m for specimen SI a smaller individual It is not known how he arrived at these numbers In 1950 he mentioned a weight of forty tonnes for the larger skeleton 23 In 1962 Edwin Harris Colbert measured a volume of 86 953 m Presuming a density of 0 9 this resulted in a weight of 78 258 kilogrammes 24 Colbert had inserted a museum model sold to the public into sand and observed the volume displaced by it Gregory S Paul in 1988 assumed that the in his opinion unrealistically high number had been caused by the fact that such models used to be very bloated compared to the real build of the animal 18 In 1980 Dale Alan Russell et al published a much lower weight of 14 8 tonnes by extrapolating from the diameter of the humerus and the thighbone 25 In 1985 the same researcher arrived at 29 tonnes by extrapolating from the circumference of these bones 26 In 1985 Robert McNeill Alexander found a value of 46 6 tonnes inserting a toy model of the British Museum of Natural History into water 27 More recent estimates based on models reconstructed from bone volume measurements which take into account the extensive weight reducing airsac systems present in sauropods and estimated muscle mass are in the range of 23 3 39 5 tonnes 25 7 43 5 short tons 17 22 In 1988 G S Paul measured a volume of 36 585 m by inserting a specially constructed model into water He estimated a weight of 31 5 tonnes assuming a low density 18 In 1994 1995 a weight of 40 tonnes extrapolating from limb bone circumference 28 In 1995 a laser scan of the skeleton was used to build a virtual model from simple geometrical shapes finding a volume of 74 42 m and concluding to a weight of 63 tonnes 29 In 2008 Gunga revised the volume using more complex shapes to 47 9 m 30 Donald Henderson in 2004 employed a computer model that calculated a volume of 32 398 m and a weight of 25 789 kilogrammes 31 Newer methods use bone wall thickness 32 However HMN SII is not the largest specimen known an assertion supported by its subadult status but HMN XV2 represented by a fibula 13 larger than the corresponding material on HMN SII 17 which might have attained 26 metres 85 ft in length 33 Gregory S Paul initially estimated the size of this specimen at 25 metres 82 ft in total length 16 metres 52 ft in total height and 45 metric tons 50 short tons in body mass 18 but later moderated at 23 metres 75 ft in total length and 40 metric tons 44 short tons in body mass 34 In 2020 Molina Perez and Larramnedi estimated the size of the HMN XV2 specimen at 25 metres 82 ft and 48 tonnes 53 short tons with a shoulder height of 6 8 metres 22 ft 2 General build edit nbsp RestorationGiraffatitan was a sauropod one of a group of four legged plant eating dinosaurs with long necks and tails and relatively small brains It had a giraffe like build with long forelimbs and a very long neck The skull had a tall arch anterior to the eyes consisting of the bony nares a number of other openings and spatulate teeth resembling chisels The first toe on its front foot and the first three toes on its hind feet were clawed Skull edit nbsp Skull cast in BerlinTraditionally the distinctive high crested skull was seen as a characteristic of the genus Brachiosaurus to which Giraffatitan brancai was originally referred however it is possible that Brachiosaurus altithorax did not show this feature since within the traditional Brachiosaurus material it is known only from Tanzanian specimens now assigned to Giraffatitan The placement of Giraffatitan nostrils has been the source of much debate with Witmer 2001 describing in Science the hypothesized position of the fleshy nostrils in Giraffatitan in as many as five possible locations Comparing the nares of dinosaurs with those of modern animals he found that all species have their external nostril openings in the front and that sauropods like Giraffatitan did not have nostrils on top of their heads but near their snouts 35 There has also been the hypothesis of various sauropods such as Giraffatitan possessing a trunk The fact that there were no narrow snouted sauropods Giraffatitan included tends to discredit such a hypothesis Stronger evidence for the absence of a trunk is found in the teeth wear of Giraffatitan which shows the kind of wear that would result from biting and tearing off of plant matter rather than purely grinding which would be the result of having already ripped the leaves and branches off with a trunk 36 Classification edit nbsp Reconstructed skeleton of the closely related BrachiosaurusWhen 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 1 Janensch concluded that the four genera must have been closely related to each other and in 1929 assigned them to a subfamily Brachiosaurinae within the family Bothrosauropodidae 37 During the twentieth century several sauropods were assigned to Brachiosauridae including Astrodon Bothriospondylus Pelorosaurus Pleurocoelus and Ultrasauros 38 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 39 40 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 41 Kingham s views of brachiosaurid taxonomy have not been accepted by many other authors 42 Since the 1990s computer based cladistic analyses allow for postulating detailed hypotheses on the relationships between species by calculating those trees 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 43 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 44 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 45 The exact status of each potential brachiosaurid varies from study to study For example a 2010 study by Chure and colleagues recognized Abydosaurus as a brachiosaurid together with Brachiosaurus which in this study included B brancai 20 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 45 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 42 while a 2013 study by Philip Mannion and colleagues had it the other way around 21 A 2012 study on titanosauriform sauropods by Michael D Emic placed Giraffatitan as sister to a clade containing Brachiosaurus and a tritomy of Abydosaurus Cedarosaurus and Venenosaurus as shown in the cladogram below 46 Brachiosauridae EuropasaurusGiraffatitan nbsp Brachiosaurus nbsp AbydosaurusCedarosaurusVenenosaurus nbsp Authentic Giraffatitan skull behind compared to that of the small EuropasaurusIn their 2024 description of Gandititan Han et al analyzed the phylogenetic relations of Macronaria focusing on titanosauriform taxa They recovered Giraffatitan as the sister taxon to Sonorasaurus similar to some previous analyses 47 in a clade also containing Brachiosaurus The results of their phylogenetic analyses are shown in the cladogram below 48 Macronaria AtlasaurusEuropasaurusGalvesaurusVouivriaLusotitanFusuisaurusTitanosauriformes SomphospondyliBrachiosauridae BrachiosaurusGiraffatitanSonorasaurusCedarosaurusVenenosaurusSoriatitanPaleobiology edit nbsp The feeding range of Giraffatitan left and SauroposeidonThe nostrils of Giraffatitan like the huge corresponding nasal openings in its skull were long thought to be located on the top of the head In past decades scientists theorized that the animal used its nostrils like a snorkel spending most of its time submerged in water in order to support its great mass The current consensus view however is that Giraffatitan was a fully terrestrial animal Studies have demonstrated that water pressure would have prevented the animal from breathing effectively while submerged and that its feet were too narrow for efficient aquatic use Furthermore new studies by Lawrence Witmer 2001 show that while the nasal openings in the skull were placed high above the eyes the nostrils would still have been close to the tip of the snout a study which also lends support to the idea that the tall crests of brachiosaurs supported some sort of fleshy resonating chamber Brain edit Giraffatitan s brain measured about 300 cubic centimetres which like those of other sauropods was small compared to its massive body size A 2009 study calculated its Encephalization Quotient a rough estimate of possible intelligence at a low 0 62 or 0 79 depending on the size estimate used Like other sauropods Giraffatitan has a sacral enlargement above the hip which some older sources misleadingly referred to as a second brain 49 However glycogen bodies are a more likely explanation 50 Nostril function edit nbsp The fleshy external nostril would have been placed at the front of the nasal fossa the depression seen in front of the bony nostrilThe bony nasal openings of neosauropods like Giraffatitan 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 51 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 52 It has been proposed that sauropods including Giraffatitan 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 53 Metabolism edit If Giraffatitan was endothermic warm blooded it would have taken an estimated ten years to reach full size if it were instead poikilothermic cold blooded then it would have required over 100 years to reach full size 54 As a warm blooded animal the daily energy demands of Giraffatitan would have been enormous it would probably have needed to eat more than 182 kg 400 lb of food per day If Giraffatitan was fully cold blooded or was a passive bulk endotherm it would have needed far less food to meet its daily energy needs Some scientists have proposed that large dinosaurs like Giraffatitan were gigantotherms 55 Internal organs of these giant sauropods were probably enormous 56 Paleoenvironment edit nbsp Restoration of two individualsGiraffatitan lived in what is now Tanzania in the Late Jurassic Tendaguru Formation 57 Since 2012 the boundary between the Kimmeridgian and Tithonian is dated at 152 1 million years ago 58 The Tendaguru ecosystem primarily consisted of three types of environment shallow lagoon like marine environments tidal flats and low coastal environments and vegetated inland environments The marine environment existed above the fair weather wave base and behind siliciclastic and ooid barriers It appeared to have had little change in salinity levels and experienced tides and storms The coastal environments consisted of brackish coastal lakes ponds and pools These environments had little vegetation and were probably visited by herbivorous dinosaurs mostly during droughts The well vegetated inlands were dominated by conifers Overall the Late Jurassic Tendaguru climate was subtropical to tropical with seasonal rains and pronounced dry periods During the Early Cretaceous the Tendaguru became more humid 59 The Tendaguru Beds are similar to the Morrison Formation of North America except in its marine interbeds 60 Giraffatitan would have coexisted with fellow sauropods like Dicraeosaurus hansemanni and D sattleri Janenschia africana Tendaguria tanzaniensis and Tornieria africanus ornithischians like Dysalotosaurus lettowvorbecki and Kentrosaurus aethiopicus the theropods Allosaurus tendagurensis Ceratosaurus roechlingi Ceratosaurus ingens Elaphrosaurus bambergi Veterupristisaurus milneri and Ostafrikasaurus crassiserratus and the pterosaur Tendaguripterus recki 61 62 63 64 Other organisms that inhabited the Tendaguru included corals echinoderms cephalopods bivalves gastropods decapods sharks neopterygian fish crocodilians and small mammals like Brancatherulum tendagurensis 65 See also edit nbsp Dinosaurs portal nbsp Wikimedia Commons has media related to Giraffatitan List of African dinosaurs Dinosaurs of Tendaguru Brachiosauridae Sauropoda SauropodomorphaReferences edit a b c d e Janensch W 1914 Ubersicht uber der Wirbeltierfauna der Tendaguru Schichten nebst einer kurzen Charakterisierung der neu aufgefuhrten Arten von Sauropoden Archiv fur Biontologie 3 1 81 110 a b Molina Perez amp Larramendi 2020 Dinosaur Facts and Figures The Sauropods and Other Sauropodomorphs New Jersey Princeton University Press p 259 Maier 2003 p 1 Maier 2003 p 3 Maier 2003 p 8 Maier 2003 p 10 Maier 2003 p 11 12 Maier 2003 p 15 Maier 2003 p 16 Maier 2003 p 17 Maier 2003 p 18 Maier 2003 p 19 Maier 2003 p 22 Maier 2003 p 23 Tamborini Marco amp Mareike Vennen 2017 Disruptions and changing habits The case of the Tendaguru expedition Museum History Journal 10 2 183 199 a b c Heinrich Wolf Dieter 1999 The Taphonomy of Dinosaurs from the Upper Jurassic of Tendaguru Tanzania Based on Field Sketches of the German Tendaguru Expedition 1909 1913 Mitteilungen aus dem Museum fur Naturkunde in Berlin Geowissenschaftliche Reihe 2 25 61 doi 10 5194 fr 2 25 1999 a b c d e f g h i j k Taylor M P 2009 A Re evaluation of Brachiosaurus altithorax Riggs 1903 Dinosauria Sauropod and its generic separation from Giraffatitan brancai Janensch 1914 PDF Journal of Vertebrate Paleontology 29 3 787 806 doi 10 1671 039 029 0309 S2CID 15220647 a b c d Paul G S 1988 The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus Giraffatitan and a comparison of the world s largest dinosaurs PDF Hunteria 2 3 1 14 Archived PDF from the original on 27 June 2022 Olshevsky G 1991 A revision of the parainfraclass Archosauria Cope 1869 excluding the advanced Crocodylia PDF Mesozoic Meanderings 2 1 196 Archived from the original PDF on 19 August 2018 Retrieved 14 April 2018 a b Chure D Britt B Whitlock J A Wilson J A 2010 First complete sauropod dinosaur skull from the Cretaceous of the Americas and the evolution of sauropod dentition Naturwissenschaften 97 4 379 391 Bibcode 2010NW 97 379C doi 10 1007 s00114 010 0650 6 PMC 2841758 PMID 20179896 a b Mannion P D Upchurch Paul Barnes Rosie N Mateus Octavio 2013 Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis Macronaria and the evolutionary history of basal titanosauriforms PDF Zoological Journal of the Linnean Society 168 98 206 doi 10 1111 zoj 12029 a b Mazzetta G V et al 2004 Giants and Bizarres Body Size of Some Southern South American Cretaceous Dinosaurs Historical Biology 16 2 4 1 13 CiteSeerX 10 1 1 694 1650 doi 10 1080 08912960410001715132 S2CID 56028251 a b Janensch W 1950 The Skeleton Reconstruction of Brachiosaurus brancai PDF Palaeontographica 97 103 Colbert E 1962 The weights of dinosaurs American Museum Novitates 2076 1 16 Russell D Beland P McIntosh J S 1980 Paleoecology of the dinosaurs of Tendaguru Tanzania Memoires de la Societe Geologique de France 59 169 175 Anderson J F Hall Martin A Russell D A 1985 Long bone circumference and weight in mammals birds and dinosaurs Journal of Zoology 207 1 53 61 doi 10 1111 j 1469 7998 1985 tb04915 x Alexander R McN 1985 Mechanics of posture and gait of some large dinosaurs Zoological Journal of the Linnean Society 83 1 1 25 doi 10 1111 j 1096 3642 1985 tb00871 x Peczkis J 1995 Implications of body mass estimates for dinosaurs Journal of Vertebrate Paleontology 14 4 520 533 doi 10 1080 02724634 1995 10011575 JSTOR 4523591 Gunga H C Kirsch K A Baartz F Rocker L Heinrich W D Lisowski W Wiedemann A Albertz J 1995 New data on the dimensions of Brachiosaurus brancai and their physiological implications Naturwissenschaften 82 4 190 192 Bibcode 1995NW 82 190G doi 10 1007 s001140050167 Gunga H C Suthau T Bellmann A Stoinski S Friedrich A Trippel T Kirsch K Hellwich O 2008 A new body mass estimation of Brachiosaurus brancai Janensch 1914 mounted and exhibited at the Museum of Natural History Berlin Germany Fossil Record 11 1 33 38 doi 10 1002 mmng 200700011 Henderson D M 2004 Tipsy Punters Sauropod Dinosaur Pneumaticity Buoyancy and Aquatic Habits Proceedings Biological Sciences 271 Supplement 4 180 183 doi 10 1098 rsbl 2003 0136 JSTOR 4142711 PMC 1810024 PMID 15252977 S2CID 23454835 Benson R B J Campione N S E Carrano M T Mannion P D Sullivan C Upchurch P Evans D C 2014 Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage PLOS Biology 12 5 e1001853 doi 10 1371 journal pbio 1001853 PMC 4011683 PMID 24802911 S2CID 16100066 Holtz Thomas R Jr 2008 Dinosaurs The Most Complete Up to Date Encyclopedia for Dinosaur Lovers of All Ages Supplementary Information Paul Gregory S 2016 The Princeton Field Guide to Dinosaurs Princeton University Press p 227 ISBN 978 1 78684 190 2 OCLC 985402380 Witmer L M 2001 Nostril position in dinosaurs and other vertebrates and its significance for nasal function PDF Science 293 5531 850 853 CiteSeerX 10 1 1 629 1744 doi 10 1126 science 1062681 PMID 11486085 S2CID 7328047 Archived from the original PDF on 6 September 2013 Naish D 20 March 2009 Junk in the trunk why sauropod dinosaurs did not possess trunks ScienceBlogs Archived from the original on 13 January 2012 Janensch 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 McIntosh John Osmolska H 1990 Sauropoda In Weishampel David B Dodson Peter Osmolska Halszka eds The Dinosauria 1 ed Berkeley University of California Press p 376 ISBN 978 0 520 06726 4 Upchurch P Barrett P M Dodson P 2004 Sauropoda In Weishampel D B Dodson P Osmolska H eds The Dinosauria Second Edition Univ of California Press Berkeley pp 259 322 ISBN 978 0 520 24209 8 Wedel Mathew J Cifelli R L Sanders R K 2000 Osteology paleobiology and relationships of the sauropod dinosaur Sauroposeidon Acta Palaeontologica Polonica 45 343 388 Kingham R F 1962 Studies of the sauropod dinosaur Astrodon Leidy Proceedings of the Washington Junior Academy of Sciences 1 38 44 a b D Emic M D 2012 The early evolution of titanosauriform sauropod dinosaurs Zoological Journal of the Linnean Society 166 3 624 671 doi 10 1111 j 1096 3642 2012 00853 x hdl 2027 42 94293 Leonardo Salgado 1993 Comments on Chubutisaurus insignis del Corro Saurischia Sauropoda Ameghiniana 30 3 265 270 Salgado L R A Coria and J O Calvo 1997 Evolution of titanosaurid sauropods I phylogenetic analysis based on the postcranial evidence Ameghiniana 34 3 32 a b Ksepka D T Norell M A 2010 The illusory evidence for Asian Brachiosauridae new material of Erketu ellisoni and a phylogenetic appraisal of basal Titanosauriformes PDF American Museum Novitates 3700 1 27 doi 10 1206 3700 2 S2CID 86254470 Archived from the original PDF on 19 August 2018 D emic Michael D 2012 The early evolution of titanosauriform sauropod dinosaurs PDF Zoological Journal of the Linnean Society 166 3 624 671 doi 10 1111 j 1096 3642 2012 00853 x hdl 2027 42 94293 S2CID 54752135 Philip D Mannion Ronan Allain Olivier Moine 2017 The earliest known titanosauriform sauropod dinosaur and the evolution of Brachiosauridae PeerJ 5 e3217 doi 10 7717 peerj 3217 PMC 5417094 PMID 28480136 Han F Yang L Lou F Sullivan C Xu X Qiu W Liu H Yu J Wu R Ke Y Xu M Hu J Lu P 2024 A new titanosaurian sauropod Gandititan cavocaudatus gen et sp nov from the Late Cretaceous of southern China Journal of Systematic Palaeontology 22 1 2293038 doi 10 1080 14772019 2023 2293038 Knoll F Schwarz Wings D 2009 Palaeoneuroanatomy of Brachiosaurus Annales de Paleontologie 95 3 165 175 doi 10 1016 j annpal 2009 06 001 Giffin Emily B 1990 Gross Spinal Anatomy and Limb Use in Living and Fossil Reptiles Paleobiology 16 4 448 485 doi 10 1017 S0094837300010186 ISSN 0094 8373 JSTOR 2400969 Witmer L M 2001 Nostril position in dinosaurs and other vertebrates and its significance for nasal function Science 293 5531 850 853 CiteSeerX 10 1 1 629 1744 doi 10 1126 science 1062681 PMID 11486085 S2CID 7328047 Czerkas S J Czerkas S A 1990 Dinosaurs a Global View Limpsfield Dragons World pp 134 135 ISBN 978 0 7924 5606 3 Knoll F Galton P M Lopez Antonanzas R 2006 Paleoneurological evidence against a proboscis in the sauropod dinosaur Diplodocus Geobios 39 2 215 221 Bibcode 2006Geobi 39 215K doi 10 1016 j geobios 2004 11 005 Case T J 1978 Speculations on the Growth Rate and Reproduction of Some Dinosaurs Paleobiology 4 3 323 doi 10 1017 S0094837300006023 Bailey J B 1997 Neural spine elongation in dinosaurs Sailbacks or buffalo backs Journal of Paleontology 71 6 1124 1146 doi 10 1017 s0022336000036076 S2CID 130861276 Jak velke vnitrni organy meli obri sauropodi 4 July 2016 Bussert Robert Heinrich Wolf Dieter Aberhan Martin 2009 The Tendaguru Formation Late Jurassic to Early Cretaceous southern Tanzania definition palaeoenvironments and sequence stratigraphy Fossil Record 12 2 141 174 doi 10 1002 mmng 200900004 Gradstein F M Ogg J G Schmitz M D amp Ogg G M 2012 A Geologic Time Scale 2012 Elsevier Aberhan Martin Bussert Robert Heinrich Wolf Dieter Schrank Eckhart Schultka Stephan Sames Benjamin Kriwet Jurgen Kapilima Saidi 2002 Palaeoecology and depositional environments of the Tendaguru Beds Late Jurassic to Early Cretaceous Tanzania Fossil Record 5 1 19 44 doi 10 1002 mmng 20020050103 Mateus Octavio 2006 Late Jurassic dinosaurs from the Morrison Formation USA the Lourinha and Alcobaca formations Portugal and the Tendaguru Beds Tanzania a comparison In Foster J R Lucas S G eds Paleontology and Geology of the Upper Morrison Formation New Mexico Museum of Natural History and Science bulletin 36 pp 223 232 ISSN 1524 4156 a href Template Cite book html title Template Cite book cite book a journal ignored help Weishampel David B et al 2004 Dinosaur distribution Late Jurassic Africa In Weishampel David B Dodson Peter and Osmolska Halszka eds The Dinosauria 2nd Berkeley University of California Press p 552 ISBN 0 520 24209 2 Barrett P M Butler R J Edwards N P amp Milner A R Pterosaur distribution in time and space an atlas p61 107 in Flugsaurier Pterosaur papers in honour of Peter Wellnhofer 2008 Hone D W E and Buffetaut E eds Zitteliana B 28 264pp 1 Rauhut Oliver W M 2011 Theropod dinosaurs from the Late Jurassic of Tendaguru Tanzania Special Papers in Palaeontology 86 195 239 Buffetaut Eric 2012 An early spinosaurid dinosaur from the Late Jurassic of Tendaguru Tanzania and the evolution of the spinosaurid dentition Oryctos 10 1 8 Heinrich Wolf Dieter et al 2001 The German Tanzanian Tendaguru Expedition 2000 Fossil Record 4 1 223 237 doi 10 1002 mmng 20010040113 Bibliography editMaier Gerhard 2003 African dinosaurs unearthed the Tendaguru expeditions Life of the Past Series ed J Farlow Indiana University Press Bloomington Indiana Retrieved from https en wikipedia org w index php title Giraffatitan amp oldid 1213700063, wikipedia, wiki, book, books, library,

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