Tornieria

Tornieria ("for Tornier") is a genus of diplodocid sauropod dinosaur from Late Jurassic of Tanzania. It has a convoluted taxonomic history.

Tornieria
Temporal range: Late Jurassic, 150 Ma
Diplodocoid skulls
Restoration of a T. africana skull (bottom right), alongside skulls of other diplodocoids.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Sauropodomorpha
Clade: Sauropoda
Family: Diplodocidae
Subfamily: Diplodocinae
Genus: Tornieria
Sternfeld, 1911
Species:
T. africana
Binomial name
Tornieria africana
Fraas, 1908 [originally Gigantosaurus]
Synonyms

Gigantosaurus africanus Fraas, 1908

Discovery and naming

500 gigantosaurus dwdu1912cropped
Historical reconstruction by Heinrich Harder (as "Gigantosaurus"), 1912

In 1907, German paleontologist Eberhard Fraas who was working the Tendaguru Beds in German East Africa (presently Tanzania), discovered two sauropod specimens at a single site ("Quarry A"). The two individuals, designated "Skeleton A" and "Skeleton B", each represented a different sauropod species. In 1908 he named these respectively Gigantosaurus africanus ("African giant lizard") and G. robustus ("Robust giant lizard").[1] A third, unrelated African species, "Gigantosaurus" dixeyi, was named by Haughton in 1928, and has since been reassigned to Malawisaurus.

However, the name Gigantosaurus had already been used for the European sauropod Gigantosaurus megalonyx Seeley, 1869. Fraas, not intending to place his species in the same genus as this English form, had believed that the name was available, since at the time the latter species was considered to be a junior synonym of Ornithopsis and Seeley in his opinion had not provided a sufficient description anyway. Another German paleontologist, Richard Sternfeld, renamed the Tanzanian sauropod Tornieria in 1911, making the two species Tornieria africana and T. robusta. The generic name honours the German herpetologist Gustav Tornier.[2]

A re-evaluation of Tornieria in 1922 by Werner Janensch concluded that one species, T. africana, was actually an African species of the North American sauropod genus Barosaurus: Barosaurus africanus.[3] The other African species, T. robusta, later turned out to belong to a titanosaur. The titanosaur species for a time was called Tornieria, but this was incorrect as T. africana had been the type species. It needed a generic name of its own and this was provided in 1991 when Rupert Wild renamed it Janenschia.

If Tornieria were the same genus as Barosaurus, then the name Tornieria would be abandoned as a junior subjective synonym. However, later researchers proposed generic distinction between the American and the African form. In the early 21st century this usage became prevalent and in 2006 Kristian Remes in a review concluded that Tornieria was indeed distinct and a valid genus.[4]

A complication is formed by the fact that Janensch in 1961 recognised a variety of B. africanus: B. africanus var. gracilis, a morph distinguished by more gracile hind limbs.[5] In 1980 John McIntosh promoted this to a full species: Barosaurus gracilis[6] which then would become a Tornieria gracilis under the present usage, a combination already published by George Olshevsky in 1992.[7] However, Remes in 2006 concluded that B. gracilis had been a nomen nudum, neither holotype nor diagnosis having been provided in 1980.[4]

Specimens and description

Elements of the original "Skeleton A" were designated by Fraas as a syntype series: SMNS 12141a, 12145a, 12143, 12140, 12142, all from the postcrania. Later some other bones from the same individual were recovered. Janensch would also refer many other fossils to B. africanus, in total 630 specimens representing at least 56 separate individuals. Of these 188 would be left after the bombardments during the Second World War. Remes, however, concluded that merely a second partial skeleton, "Skeleton k", including also some skull elements, could be reliably referred, and a series of caudal vertebrae. The remains are from the later strata of the Tendaguru, the obere Dinosauriermergel or "Upper Dinosaur Marl", dating from the Tithonian.[4]

Tornieria was a large sauropod, with a maximum known femur length of 138 centimetres (54 in) suggesting an animal around the same size as Barosaurus; 26 m (85 ft) and 23 metric tons.[8] It shared elongated neck vertebrae and a rather long forelimb with Barosaurus. However, it differed from the American form by details in the anterior caudal vertebrae and from Barosaurus and Diplodocus both by its plesiomorphic hindlimb proportions with a short lower leg.

Phylogeny

After performing a cladistic analysis, Remes (2006) concluded that Tornieria was the sister taxon of a clade formed by Barosaurus and Diplodocus. It would thus be a member of the Diplodocinae.[4] The following cladogram is based on the phylogenetic analysis conducted by Whitlock in 2011, showing the relationships of Tornieria among the other genera assigned to the taxon Diplodocidae:[9]

Diplodocidae

Apatosaurus

Supersaurus

Dinheirosaurus

Tornieria

Barosaurus

Diplodocus

References

  1. ^ E. Fraas, 1908, "Dinosaurierfunde in Ostafrika", Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg 64: 84-86
  2. ^ Sternfeld, R., 1911, "Zur Nomenklatur der Gattung Gigantosaurus Fraas", Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin 1911: 398
  3. ^ Janensch, W., 1922, "Das Handskelett von Gigantosaurus robustus und Brachiosaurus brancai aus den Tendaguru-Schichten Deutsch- Ostafrikas", Centralblatt für Mineralogie, Geologie und Paläontologie 1922: 464–480
  4. ^ a b c d Remes, K., 2006, "Revision of the Tendaguru sauropod Tornieria africana (Fraas) and its relevance for sauropod paleobiogeography", Journal of Vertebrate Paleontology 26(3): 651–669
  5. ^ Janensch, W., 1961, "Die Gliedmaßen und Gliedmaßengürtel der Sauropoden der Tendaguru-Schichten", Palaeontographica Supplement 7(3): 177–235
  6. ^ D. Russell, P. Beland, and J. McIntosh, 1980, "Paleoecology of the dinosaurs of Tendaguru (Tanzania)", Memoires de la Societé Geologique de France, N.S. 139: 169-175
  7. ^ G. Olshevsky, 1992, A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings 2, pp. 1-268
  8. ^ "Giraffatitan: Your Dinosaurs Are Wrong #18". YouTube. Retrieved 28 January 2019.
  9. ^ Whitlock, J.A. (2011). "A phylogenetic analysis of Diplodocoidea (Saurischia: Sauropoda)." Zoological Journal of the Linnean Society, Article first published online: 12 Jan 2011.
Australodocus

Australodocus (meaning "southern beam" from the Latin australis "southern" and the Greek dokos/δοκоς "beam") is a genus of sauropod dinosaur that lived during the Late Jurassic period, around 150 million years ago, in what is now Tanzania. Though initially considered a diplodocid, recent analyses suggest it may instead be a titanosauriform.

Barosaurus

Barosaurus ( BARR-o-SAWR-əs) was a giant, long-tailed, long-necked, plant-eating dinosaur closely related to the more familiar Diplodocus. Remains have been found in the Morrison Formation from the Upper Jurassic Period of Utah and South Dakota (and possibly Africa, as exemplified by the Kadsi Formation). It is present in stratigraphic zones 2-5.The composite term Barosaurus comes from the Greek words barys (βαρυς) meaning "heavy" and sauros (σαυρος) meaning "lizard"; thus "heavy lizard".

Dinheirosaurus

Dinheirosaurus is a genus of diplodocid sauropod dinosaur that is known from fossils uncovered in modern-day Portugal. It may represent a species of Supersaurus. The only species is Dinheirosaurus lourinhanensis, first described by José Bonaparte and Octávio Mateus in 1999 for vertebrae and some other material from the Lourinhã Formation. Although the precise age of the formation is not known, it can be dated around the early Tithonian of the Late Jurassic.

The known material includes two cervical vertebrae, nine dorsal vertebrae, a few ribs, a fragment of a pubis, and many gastroliths. Of the material, only the vertebrae are diagnostic, with the ribs and pubis being too fragmentary or general to distinguish Dinheirosaurus. This material was first described as in the genus Lourinhasaurus, but differences were noticed and in 1999 Bonaparte and Mateus redescribed the material under the new binomial Dinheirosaurus lourinhanensis. Another specimen, ML 418, thought to be Dinheirosaurus, is now known to be from another Portuguese diplodocid. This means that Dinheirosaurus lived alongside many theropods, sauropods, thyreophorans and ornithopods, as well as at least one other diplodocid.

Dinheirosaurus is a diplodocid, a relative of Apatosaurus, Diplodocus, Barosaurus, Supersaurus, and Tornieria. Among those, the closest relative to Dinheirosaurus is Supersaurus.

Diplodocidae

Diplodocids, or members of the family Diplodocidae ("double beams"), are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the Earth, including Diplodocus and Supersaurus, some of which may have reached lengths of up to 34 metres (112 ft).

Diplodocinae

Diplodocinae is an extinct subfamily of diplodocid sauropods that existed from the Late Jurassic to Early Cretaceous of North America, Europe and South America, about 161.2 to 136.4 million years ago. Genera within the subfamily include Tornieria, Supersaurus, Leinkupal, Galeamopus, Diplodocus, Kaatedocus and Barosaurus.Cladogram of the Diplodocidae after Tschopp, Mateus, and Benson (2015).

Diplodocoidea

Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like Supersaurus, Diplodocus, Apatosaurus, and Amphicoelias. Most had very long necks and long, whip-like tails; however, one family (the dicraeosaurids) are the only known sauropods to have re-evolved a short neck, presumably an adaptation for feeding low to the ground. This adaptation was taken to the extreme in the highly specialized sauropod Brachytrachelopan. A study of snout shape and dental microwear in diplodocoids showed that the square snouts, large proportion of pits, and fine subparallel scratches in Apatosaurus, Diplodocus, Nigersaurus, and Rebbachisaurus suggest ground-height nonselective browsing; the narrow snouts of Dicraeosaurus, Suuwassea, and Tornieria and the coarse scratches and gouges on the teeth of Dicraeosaurus suggest mid-height selective browsing in those taxa. This taxon is also noteworthy because diplodocoid sauropods had the highest tooth replacement rates of any vertebrates, as exemplified by Nigersaurus, which had new teeth erupting every 30 days.

Gigantosaurus

Gigantosaurus (from the Greek "Γίγας" and "σαυρος", meaning "giant lizard") is a sauropod dinosaur genus from the Late Jurassic Kimmeridge Clay Formation of England. The type species, Gigantosaurus megalonyx, was named and described by Harry Govier Seeley in 1869. Its syntype series consists of several separately discovered sauropod bones found in Cambridgeshire, including two caudal (tail) vertebrae (CAMSM J.29477 and CAMSM J.29478), the distal end of a tibia (CAMSM J.29483), a cast of the right radius (CAMSM J.29482), a cast of phalanx (CAMSM J.29479) and an osteoderm (CAMSM J.29481). It was synonymised to Ornithopsis humerocristatus by Richard Lydekker in 1888 and to Pelorosaurus by Friedrich von Huene in 1909. Today it is considered a nomen dubium.

Because of these references Eberhard Fraas incorrectly assumed in 1908 the name was available for other species and he used it, despite it being preoccupied, for African material totally unrelated to the British finds. As a result, the name Gigantosaurus factored into the convoluted taxonomic history of the African dinosaurs Barosaurus, Tornieria, and Janenschia. A discussion of this can be found in the main Tornieria article.

Gravisauria

Gravisauria is a clade of sauropod dinosaurs consisting of some genera, Vulcanodontidae and Eusauropoda.

Janenschia

Janenschia (named after Werner Janensch) is a large herbivorous sauropod dinosaur from the Late Jurassic Tendaguru Formation of Tanzania, Africa, 155 million years ago.

Kaatedocus

Kaatedocus is a genus of diplodocine flagellicaudatan sauropod known from the middle Late Jurassic (Kimmeridgian stage) of northern Wyoming, United States. It is known from well-preserved skull and cervical vertebrae which were collected in the lower part of the Morrison Formation. The type and only species is Kaatedocus siberi, described in 2012 by Emanuel Tschopp and Octávio Mateus.

Kaijutitan

Kaijutitan (meaning "Kaiju titan" after the type of Japanese movie monsters) is a genus of basal titanosaur dinosaur from the Sierra Barrosa Formation from Neuquén Province in Argentina. The type and only species is Kaijutitan maui.

Leinkupal

Leinkupal is a genus of diplodocine sauropod known from the Early Cretaceous (Late Berriasian to Early Valanginian stage) of the Bajada Colorada Formation, southeastern Neuquén Basin in the Neuquén Province of Argentina. It contains a single species, Leinkupal laticauda.

List of African dinosaurs

This is a list of dinosaurs whose remains have been recovered from Africa. Africa has a rich fossil record, but it is patchy and incomplete. It is rich in Triassic and Early Jurassic dinosaurs. African dinosaurs from these time periods include Coelophysis, Dracovenator, Melanorosaurus, Massospondylus, Euskelosaurus, Heterodontosaurus, Abrictosaurus, and Lesothosaurus. In the Middle Jurassic, the sauropods Atlasaurus, Chebsaurus, Jobaria, and Spinophorosaurus, flourished, as well as the theropod Afrovenator. The Late Jurassic is well represented in Africa, mainly thanks to the spectacular Tendaguru Formation. Veterupristisaurus, Ostafrikasaurus, Elaphrosaurus, Giraffatitan, Dicraeosaurus, Janenschia, Tornieria, Tendaguria, Kentrosaurus, and Dysalotosaurus are among the dinosaurs whose remains have been recovered from Tendaguru. This fauna seems to show strong similarities to that of the Morrison Formation in the United States and the Lourinha Formation in Portugal. For example, similar theropods, ornithopods and sauropods have been found in both the Tendaguru and the Morrison. This has important biogeographical implications.

The Early Cretaceous in Africa is known primarily from the northern part of the continent, particularly Niger. Suchomimus, Elrhazosaurus, Rebbachisaurus, Nigersaurus, Kryptops, Nqwebasaurus, and Paranthodon are some of the Early Cretaceous dinosaurs known from Africa. The Early Cretaceous was an important time for the dinosaurs of Africa because it was when Africa finally separated from South America, forming the South Atlantic Ocean. This was an important event because now the dinosaurs of Africa started developing endemism because of isolation.

The Late Cretaceous of Africa is known mainly from North Africa. During the early part of the Late Cretaceous, North Africa was home to a rich dinosaur fauna. It includes Spinosaurus, Carcharodontosaurus, Rugops, Bahariasaurus, Deltadromeus, Paralititan, Aegyptosaurus, and Ouranosaurus.

Lithostrotia

Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Unchurch et al. in 2004 as the most recent common ancestor of Malawisaurus and Saltasaurus and all the descendants of that ancestor. Lithostrotia is derived from the Ancient Greek lithostros, meaning "inlaid with stones", referring to the fact that many known lithostrotians are preserved with osteoderms. However, osteoderms are not a distinguishing feature of the group, as the two noted by Unchurch et al. include caudal vertebrae with strongly concave front faces (procoely), although the farthest vertebrae are not procoelous.

Malawisaurus

Malawisaurus (meaning "Malawi lizard") was a genus of sauropod dinosaur (specifically a titanosaurian). It lived in what is now Africa, specifically Malawi, during the Aptian age of the Early Cretaceous Period. It is one of the few titanosaurs for which skull material has been found.

T. africana

T. africana may refer to:

Tabernaemontana africana, a flowering plant species in the genus Tabernaemontana

Tieghemella africana, a plant species found in Cameroon, the Republic of the Congo and Gabon

Tornieria africana, a dinosaur species

Treculia africana, the African breadfruit, a tree species

Tricalysia africana, a plant species endemic to South Africa

T. gracilis

T. gracilis may refer to:

Taterillus gracilis, the gracile tateril or slender gerbil, a rodent species found in Africa

Thomasomys gracilis, the slender Oldfield mouse, a rodent species found in Ecuador and Peru

Tornieria gracilis, a dinosaur species

Tribolonotus gracilis, a skink species found in New Guinea

Trimeresurus gracilis, a venomous pitviper species found only in Taiwan

Troglohyphantes gracilis, a Kočevje subterranean spider species

Trypeta gracilis, a fruit fly species

Tupaia gracilis, the slender treeshrew, a treeshrew species found in Indonesia and Malaysia

Thalassina gracilis, a species of mud lobster in the genus Thalassina

Tambatitanis

Tambatitanis is an extinct genus of titanosauriform dinosaur from the Early Cretaceous (probably early Albian) of Japan. It is known from a single type species, Tambatitanis amicitiae. It was probably around 14 meters long and its mass was estimated at some 4 tonnes. It was a basal titanosauriform and possibly belonged to the Euhelopodidae.

Wamweracaudia

Wamweracaudia is a large herbivorous sauropod dinosaur from the Late Jurassic Tendaguru Formation of Tanzania, Africa, 155 million years ago.

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