Sauropodomorpha (/ˌsɔːrəˌpɒdəˈmɔːrfə/ SOR-ə-POD-ə-MOR-fə; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The "prosauropods", which preceded the sauropods, were smaller and were often able to walk on two legs. The sauropodomorphs were the dominant terrestrial herbivores throughout much of the Mesozoic Era, from their origins in the mid-Triassic (approximately 230 Ma) until their decline and extinction at the end of the Cretaceous (approximately 66 Ma).
|Mounted skeleton of Plateosaurus engelhardti at Eberhard-Karls-University|
Sauropodomorphs were adapted to browsing higher than any other contemporary herbivore, giving them access to high tree foliage. This feeding strategy is supported by many of their defining characteristics, such as: a light, tiny skull on the end of a long neck (with ten or more elongated cervical vertebrae) and a counterbalancing long tail (with one to three extra sacral vertebrae).
Their teeth were weak, and shaped like leaves or spoons (lanceolate or spatulate). Instead of grinding teeth, they had stomach stones (gastroliths), similar to the gizzard stones of modern birds and crocodiles, to help digest tough plant fibers. The front of the upper mouth bends down in what may be a beak.
One of the earliest known sauropodomorphs, Saturnalia, was small and slender (1.5 metres, or 5 feet long); but, by the end of the Triassic, they were the largest dinosaurs of their time, and throughout the Jurassic and Cretaceous they kept on growing. Ultimately the largest sauropods, like Supersaurus, Diplodocus hallorum, Patagotitan, and Argentinosaurus, reached 30–40 metres (98–131 ft) in length, and 60,000–100,000 kilograms (65–110 US short tons) or more in mass.
Initially bipedal, as their size increased they evolved a four-legged graviportal gait adapted only to walking slowly on land, like elephants. The early sauropodomorphs were most likely omnivores as their shared common ancestor with the other saurischian lineage (the theropods) was a carnivore. Therefore, their evolution to herbivory went hand in hand with their increasing size and neck length.
Among the very first dinosaurs to evolve in the Late Triassic Period, about 230 million years ago (Mya), they became the dominant herbivores by halfway through the late Triassic (during the Norian stage). Their perceived decline in the early Cretaceous is most likely a bias in fossil sampling, as most fossils are known from Europe and North America. Sauropods were still the dominant herbivores in the Gondwanan landmasses, however. The spread of flowering plants (angiosperms) and "advanced" ornithischians, another major group of herbivorous dinosaurs (noted for their highly developed chewing mechanisms), are most likely not a major factor in sauropod decline in the northern continents. Like all non-avian dinosaurs, the sauropodomorphs became extinct 66 Mya, during the Cretaceous–Paleogene extinction event.
The earliest and most basal sauropodomorphs known are Chromogisaurus novasi and Panphagia protos, both from the Ischigualasto Formation, dated to 231.4 million years ago (late Ladinian age of the Middle Triassic according to the ICS; alternately called the early Carnian age of the Late Triassic in the system used by the Geological Society of America). Some studies have found Eoraptor lunensis (also from the Ischigualasto Formation), traditionally considered a theropod, to be an early member of the sauropodomorph lineage, which would make it the most basal sauropodomorph known.
Sauropodomorpha is one of the two major clades within the order Saurischia. The sauropodomorphs' sister group, the Theropoda, includes bipedal carnivores like Velociraptor and Tyrannosaurus; as well as birds. However, sauropodomorphs also share a number of characteristics with the Ornithischia, so a small minority of palaeontologists, like Bakker, have historically placed both sets of herbivores within a group called "Phytodinosauria" or "Ornithischiformes".
In Linnaean taxonomy, Sauropodomorpha (which means "lizard feet forms") is either a suborder or is left unranked. It was originally established by Friedrich von Huene in 1932, who broke it into two groups: the basal forms within Prosauropoda, and their descendants, the giant Sauropoda.
Phylogenetic analyses by Adam Yates (2004, 2006) and others firmly placed Sauropoda within a paraphyletic "Prosauropoda". Recent cladistic analyses suggest that the clade Prosauropoda, which was named by Huene in 1920 and was defined by Sereno, in 1998, as all animals more closely related to Plateosaurus engelhardti than to Saltasaurus loricatus, is a junior synonym of Plateosauridae as both contain the same taxa.
Most modern classification schemes break the prosauropods into a half-dozen groups that evolved separately from one common lineage. While they have a number of shared characteristics, the evolutionary requirements for giraffe-like browsing high in the trees may have caused convergent evolution, where similar traits evolve separately because they faced the same evolutionary pressure, instead of (homologous) traits derived from a shared ancestor.
Below is a cladogram of basal sauropodomorpha after Otero et al., 2015.
A large phylogenetic analysis of early dinosaurs published by Matthew Baron, David Norman and Paul Barrett (2017) in the journal Nature redefined Sauropodomorpha and Saurischia and recovered Herrerasauridae as the sister group to Sauropodomorpha within Saurischia. This resulted from the proposed removal of Theropoda from Saurischia and the formation of Ornithoscelida, a clade containing Theropoda and Ornithischia.
Comparisons between the scleral rings of several sauropodmorph genera (Diplodocus, Lufengosaurus, Nemegtosaurus, Plateosaurus, and Riojasaurus) and modern birds and reptiles suggest that they may have been cathemeral, active throughout the day at short intervals.
Sauropodomorphs reached the age of sexual maturity well before they were fully-grown adults. A study by Griebeler et al. (2013) concluded that the maximum growth rates of sauropodomorphs were comparable to those of precocial birds and the black rhinoceros but lower than the growth rates of average mammals.
A long-standing hypothesis has been that early sauropodomorphs were carnivorous, as expected for most early dinosaurs. This hypothesis is supported by the current basalmost sauropodomorph, Buriolestes, and its plesiomorphic, theropod-like teeth. The teeth of more derived sauropodomorphs such as Eoraptor, Panphagia, and Pampadromaeus are better-suited for herbivorous (or possibly omnivorous) diets. Regardless of the phylogenetic position of herrerasaurids or Eoraptor (which are in flux), ancestral state reconstructions recover carnivory as being ancestral to sauropodomorphs.
Adeopapposaurus (meaning "far eating lizard", in reference to its long neck) is a genus of prosauropod dinosaur from the Early Jurassic Cañón del Colorado Formation of San Juan, Argentina. It was similar to Massospondylus. Four partial skeletons with two partial skulls are known.The type specimen, PVSJ568, includes a skull and most of a skeleton to just past the hips. The form of the bones at the tips of the upper and lower jaws suggests it had keratinous beaks. The fossils now named Adeopapposaurus were first thought to represent South American examples of Massospondylus; while this is no longer the case, Adeopapposaurus is classified as a massospondylid. Adeopapposaurus was described in 2009 by Ricardo N. Martínez. The type species is A. mognai, referring to the Mogna locality where it was found.Anchisauria
The Anchisauria were a clade of sauropodomorph dinosaurs that lived during the Late Triassic and Early Jurassic. The name Anchisauria was first used by Galton and Upchurch in the second edition of The Dinosauria. Galton and Upchurch assigned two families of dinosaurs to the Anchisauria: the Anchisauridae and the Melanorosauridae. The more common prosauropods Plateosaurus and Massospondylus were placed in the sister clade Plateosauria.
However, recent research indicates that Anchisaurus is closer to sauropods than traditional prosauropods; thus, Anchisauria would also include Sauropoda.The following cladogram simplified after an analysis presented by Blair McPhee and colleagues in 2014.Chromogisaurus
Chromogisaurus is a sauropodomorph which existed in Argentina during the Late Triassic (Carnian) period. It was a herbivore about 2 metres (6 ft 7 in) in length, and was optionally quadrupedal.Euskelosaurus
Euskelosaurus browni ("good leg lizard") is a semi-bipedal plateosaurid sauropodomorph dinosaur from the early Triassic of South Africa and Lesotho. Fossils have only been recovered from the lower Elliot Formation in South Africa and Lesotho, and in one locality in Zimbabwe.Guaibasauridae
Guaibasauridae is a family of basal saurischian dinosaurs, known from fossil remains of late Triassic period formations in Brazil and Argentina.Jingshanosaurus
Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.Melanorosauridae
The Melanorosauridae were a family of sauropodomorph dinosaurs which lived during the Late Triassic and Early Jurassic. The name Melanorosauridae was first coined by Friedrich von Huene in 1929. Huene assigned several families of dinosaurs to the infraorder "Prosauropoda": the Anchisauridae, the Plateosauridae, the Thecodontosauridae, and the Melanorosauridae. Since then, these families have undergone numerous revisions. Galton and Upchurch (2004) considered Camelotia, Lessemsaurus, and Melanorosaurus members of the family Melanorosauridae. A more recent study by Yates (2007) indicates that the melanorosaurids were instead early sauropods.Ngwevu
Ngwevu intloko (pronounced 'Ng-g'where-voo; directly from Xhosa 'ngwevu' and 'intloko' meaning "grey skull") is a genus of massospondylid sauropodomorph dinosaur from the Lower Jurassic Elliot Formation of South Africa. The type and only known specimen, BP/1/4779, was discovered in 1978 by James William Kitching. It had in 1990 and 2004 been regarded as an unusual specimen of the related Massospondylus, with a horizontally and vertically compressed skull, but in 2019 the specimen was after restudy concluded to belong to a new distinct genus. The genus is primarily distinguished by its skull being more robust than that of Massospondylus.Plateosauria
Plateosauria is a clade of sauropodomorph dinosaurs which lived during the Late Triassic to the Late Cretaceous. The name Plateosauria was first coined by Gustav Tornier in 1913. The name afterwards fell out of use until the 1980s.
Plateosauria is a node-based taxon. In 1998, Paul Sereno defined Plateosauria as the last common ancestor of Plateosaurus engelhardti and Massospondylus carinatus, and its descendants. Peter Galton and Paul Upchurch in 2004 used a different definition: the last common ancestor of Plateosaurus engelhardti and Jingshanosaurus xinwaensis, and its descendants. In their cladistic analysis the Plateosauria belonged to the Prosauropoda, and included the Plateosauridae subgroup. In Galton's and Upchurch's study also Coloradisaurus, Euskelosaurus, Jingshanosaurus, Massospondylus, Mussaurus, Sellosaurus, and Yunnanosaurus proved to be plateosaurians.However, recent cladistic analyses suggest that the Prosauropoda as traditionally defined is paraphyletic to sauropods. Prosauropoda, as currently defined, is a synonym of Plateosauridae as both contain the same taxa by definition.
The following cladogram simplified after an analysis presented by Apaldetti and colleagues in 2011.
The following cladogram simplified after an analysis presented by Blair McPhee and colleagues in 2014.Plateosauridae
Plateosauridae is a family of plateosaurian sauropodomorphs from the Late Triassic of Europe. Although several dinosaurs have been classified as plateosaurids over the years, the family Plateosauridae is now restricted to Plateosaurus. In another study, Yates (2003) sunk Sellosaurus into Plateosaurus (as P. gracilis).Pulanesaura
Pulanesaura is an extinct genus of basal sauropod known from the Early Jurassic (late Hettangian to Sinemurian) Upper Elliot Formation of the Free State, South Africa. It contains a single species, Pulanesaura eocollum, known from partial remains of at least two subadult to adult individuals.Riojasauridae
Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).Riojasaurus
Riojasaurus (meaning "Rioja lizard") was a herbivorous sauropodomorph dinosaur named after La Rioja Province in Argentina where it was found in the Los Colorados Formation in the Ischigualasto-Villa Unión Basin by José Bonaparte. It lived during the Late Triassic (Norian stage) and grew to about 10 metres (33 ft) long. Riojasaurus is the only known riojasaurid to live in South America.Ruehleia
Ruehleia is a genus of sauropodomorph dinosaur from the Late Triassic period of Germany. The type species is R. bedheimensis, described by Galton in 2001, and is named for the German paleontologist Hugo Ruehle von Lilienstern. The fossils consist of one nearly complete skeleton, consisting of cervical (neck), dorsal (back), and caudal (tail) vertebrae; a partial sacrum; a scapula-coracoid; pelvic bones; most of the limb bones; and partially complete manus (hands).
The fossils were found in central Germany and date to the Norian stage, around 216 to 203 million years ago.Saturnaliinae
Saturnaliinae is a clade of sauropodomorph dinosaurs found in Brazil and Argentina.
In 2010, Martin Ezcurra defined the subfamily Saturnaliinae for the clade containing Saturnalia and Chromogisaurus, which were found to be close relatives in several studies. While they are sometimes found to be a subgroup within the Guaibasauridae, other studies have found the saturnaliines to form an independent lineage at the very base of the sauropodomorph family tree. Langer and colleagues (2019) recovered Pampadromaeus and Panphagia as relatives of Saturnalia and Chromogisaurus, elevating Saturnaliinae to family rank as Saturnaliidae. They recovered Guaibasaurus as a basal theropod.Unaysauridae
Unaysauridae is a family of basal sauropodomorphs from the Late Triassic of India and Brazil.Xixiposaurus
Xixiposaurus is a genus of prosauropod dinosaur which existed in what is now Lower Lufeng Formation, China during the lower Jurassic period. It was first named by Sekiya Toru in 2010 and the type species is Xixiposaurus suni.