Giraffatitan

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]

The quarries listed above represent only the most important sites where Giraffatitan 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.

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.

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

In 1988, Gregory S. Paul noted that Brachiosaurus brancai (on which most popular depictions of Brachiosaurus were based) showed significant differences from the North American Brachiosaurus, especially in the proportions of its trunk vertebrae and in its more gracile build. Paul used these differences to create a subgenus he named Brachiosaurus (Giraffatitan) brancai. In 1991, George Olshevsky asserted that these differences were enough to place the African brachiosaurid in its own genus, simply Giraffatitan.[34]

Further differences between the African and North American forms came to light with the description in 1998 of a North American Brachiosaurus skull. This skull, which had been found nearly a century earlier (it is the skull Marsh used on his early reconstructions of Brontosaurus), is identified as "Brachiosaurus sp." and may well belong to B. altithorax. The skull is closer to Camarasaurus in some features such as the form of the front teeth and more elongated and less hollowed-out on top than the distinctive short-snouted and high-crested skull of Giraffatitan.[35]

The classification of Giraffatitan as a separate genus was not widely followed by other scientists at first, as it was not supported by a rigorous comparison of both species. However, a detailed comparison was published by Michael P. Taylor in 2009. Taylor showed that "Brachiosaurus" brancai differed from B. altithorax in almost every fossil bone that could be compared, in terms of both size, shape, and proportion, concluding that the placement of Giraffatitan in a separate genus was valid.[17] Taylor found evidence of a sister relationship between Giraffatitan and Brachiosaurus, although his analysis omitted other Brachiosaurids.[17] A more recent study on Titanosauriform sauropods by D'Emic (2012)[36] places Giraffatitan as sister to a clade containing Brachiosaurus and a tritomy of Abydosaurus, Cedarosaurus, and Venenosaurus as shown below:

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).

Restoration of two individuals

Giraffatitan lived in what is now Tanzania in the Late Jurassic Tendaguru Formation.[42] Since 2012, the boundary between the Kimmeridgian and Tithonian is dated at 152.1 million years ago.[43]

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.[44] The Tendaguru Beds are similar to the Morrison Formation of North America except in its marine interbeds.[45]

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.[46][47][48][49] Other organisms that inhabited the Tendaguru included corals, echinoderms, cephalopods, bivalves, gastropods, decapods, sharks, neopterygian fish, crocodilians and small mammals like Brancatherulum tendagurensis.[50]

• List of African dinosaurs
• Dinosaurs of Tendaguru
• Brachiosauridae
• Sauropoda
• Sauropodomorpha

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

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