Camarasaurid
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Camarasauridae is a family of
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
. Among sauropods, camarasaurids are small to medium-sized, with relatively short necks. They are visually identifiable by a short skull with large nares, and broad, spatulate teeth filling a thick jaw.1. Weishampel, D. B., Dodson, P., & Osmólska, H. (Eds.). (1990). ''The dinosauria''. Univ of California Press. Based on cervical vertebrae and cervical rib biomechanics, camarasaurids most likely moved their necks in a vertical, rather than horizontal, sweeping motion, in contrast to most diplodocids.


Taxonomy

Camarasauridae was named by
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
in 1877. Its
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
is ''
Camarasaurus ''Camarasaurus'' ( ) is a genus of sauropod dinosaur that lived in North America during the Late Jurassic period. Its fossil remains have been found in the Morrison Formation, dating to the Kimmeridgian and Tithonian ages of the Jurassic, betwe ...
'', and it is defined as the clade containing all species more closely related to ''
Camarasaurus supremus ''Camarasaurus'' ( ) is a genus of sauropod dinosaur that lived in North America during the Late Jurassic period. Its fossil remains have been found in the Morrison Formation, dating to the Kimmeridgian and Tithonian ages of the Jurassic, betwee ...
'' than '' Saltasaurus loricatus''.


Phylogenetic relationships

Camarasauridae is typically regarded as belonging to
Macronaria Macronaria is a clade of sauropod dinosaurs. Macronarians are named after the large diameter of the nasal opening of their skull, known as the external naris, which exceeded the size of the orbit, the skull opening where the eye is located (hence ...
, one of the two major branches of
Neosauropoda Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentina, Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recent ...
. Within Macronaria, it occupies a basal position, outside of
Titanosauriformes Macronaria is a clade of sauropod dinosaurs. Macronarians are named after the large diameter of the nasal opening of their skull, known as the external naris, which exceeded the size of the orbit, the skull opening where the eye is located (hence ...
. However, some studies have found Camarasauridae to lie outside Neosauropoda.


Members

''Camarasaurus'' is the only taxon uncontroversially regarded as a valid genus of camarasaurid. It contains four species: '' C. grandis'', '' C. lentus'', '' C. lewisi'', and '' C. supremus''. ''C. lewisi'' may represent a distinct genus, ''Cathetosaurus''. ''
Lourinhasaurus ''Lourinhasaurus'' is an extinct genus of herbivorous sauropod dinosaur genus dating from Late Jurassic strata of Estremadura, Portugal. The genus is monotypic, containing one species, ''Lourinhasaurus alenquerensis''. The type specimen for thi ...
'', the type species of which was formerly assigned to ''Camarasaurus'', is regarded as a camarasaurid by most studies, though it has also been considered to be a basal eusauropod. ''
Oplosaurus ''Oplosaurus'' (meaning "armed or weapon lizard" or "armoured lizard"; see below for discussion) was a genus of sauropod dinosaur from the Barremian-age Lower Cretaceous Wessex Formation of the Isle of Wight, England. It is known from a single to ...
'', from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
of the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...
, has been suggested to be a camarasaurid, but as it is only known from a tooth, its position within Eusauropoda is difficult to determine. '' Tehuelchesaurus'', from the Late Jurassic of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, has been considered a camarasaurid in some studies, but a wide range of other phylogenetic positions have been proposed, including a close relationship to ''
Omeisaurus ''Omeisaurus'' (meaning "Omei lizard") is a genus of sauropod dinosaur from the Middle to Late Jurassic Period (Bathonian- Oxfordian stage) of what is now China. Its name comes from Mount Emei, where it was discovered in the lower Shaximiao ...
'', a position in
Turiasauria Turiasauria is an unranked clade of eusauropod dinosaurs known from Middle Jurassic to Early Cretaceous deposits in Europe, North America, and Africa. Description Turiasauria was originally erected by Royo-Torres et al. (2006) to include '' Tur ...
, or as a non-camarasaurid basal macronarian. ''
Bellusaurus ''Bellusaurus'' (meaning "Beautiful lizard", from Vulgar Latin ''bellus'' 'beautiful' (masculine form) and Ancient Greek ''sauros'' 'lizard') was a sauropod dinosaur from the Late Jurassic ( Oxfordian) known from juvenile specimens that would hav ...
'', which is only known from juvenile remains, may be a camarasaurid, though it has also been considered a basal macronarian, turiasaur, or mamenchisaurid.


Former members

In 1970, Rodney Steel took an expansive concept of Camarasauridae, encompassing all sauropods then known except diplodocoids and titanosaurs. In 1990, John S. McIntosh regarded Camarasauridae as made up of two subfamilies: Camarasaurinae, containing ''Camarasaurus'', ''Aragosaurus'', ''Euhelopus'', and ''Tienshanosaurus'', and Opisthocoelicaudiinae, containing ''Opisthocoelicaudia'' and ''Chondrosteosaurus''. ''
Dashanpusaurus ''Dashanpusaurus'' (meaning "Dashanpu lizard" after the township it was discovered in) is an extinct genus of sauropod dinosaur that lived during the middle of the Jurassic period. The dinosaur was described in 2005 by Peng Guangzhao, Ye Yong, Ga ...
'', from the Middle Jurassic of China, was originally described as a camarasaurid, but was subsequently found to be a basal macronarian by phylogenetic analysis.


Diagnostic characters

Several skeletal features have been used to characterize the camarasaurids. In the skull, these include an external narial diameter approximately 40% of the long-axis length of the skull, an arched internarial bar, a short muzzle anterior to the nares, and maxillary shelf. In the rest of the axial skeleton, these include flat ventral faces on the cervical vertebrae, a triangular flare to the neural spines of the middle and posterior dorsal vertebrae, and a concave posterior surface to the anterior thoracic ribs, as well as an external haemal canal across the anterior vertebrae of the tail.Wilson, J. A., & Sereno, P. C. (1998). Early evolution and higher-level phylogeny of sauropod dinosaurs. ''Journal of Vertebrate Paleontology'', ''18''(S2), 1-79. A reduction to two carpals, long metacarpals relative to the radius, and a twisted ischial shaft serve to identify the appendicular skeleton.


Palaeobiogeography

Broadly speaking, camarasaurids occupied a distribution limited to the
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
n continent during the
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
. Most currently accepted camarasaurid specimens have been discovered in the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
of North America, however some specimens from the African
Tendaguru Formation The Tendaguru Formation, or Tendaguru Beds are a highly List of fossiliferous stratigraphic units in Tanzania, fossiliferous Lithostratigraphy, formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represe ...
have been speculated to belong to the genus ''Camarasaurus'', and the closely related ''Lourinhasaurus'' was found in Portugal.Raath, J. S. (1987). Sauropod dinosaurs from the Central Zambezi Valley, Zimbabwe, and the age of the Kadzi Formation. ''South African Journal of Geology'', ''90''(2), 107-119.


References

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