Eusauropod
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Eusauropoda (meaning "True Lizard Foot") is a derived
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
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 ...
dinosaur 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 ...
s. Eusauropods represent the node-based group that includes all descendant sauropods starting with the basal eusauropods of ''
Shunosaurus ''Shunosaurus'', meaning "Lizard from Sichuan", is a genus of sauropod dinosaur from Late Jurassic ( Oxfordian) beds in Sichuan Province in China, from 161 to 157 Million years ago. The name derives from "Shu", an ancient name for the Sichuan pr ...
'', and possibly ''
Barapasaurus ''Barapasaurus'' ( ) is a genus of Basal (phylogenetics), basal sauropod dinosaur from Jurassic rocks of India. The only species is ''B. tagorei''. ''Barapasaurus'' comes from the lower part of the Kota Formation, which is of Early to Middle J ...
'', and ''
Amygdalodon ''Amygdalodon'' (; "almond tooth" for its almond shaped teeth) was a genus of basal sauropod from the Middle Jurassic of Argentina. The type species is ''Amygdalodon patagonicus''. Fossils of ''Amygdalodon'' have been found in the Toarcian Cerro ...
'', but excluding ''
Vulcanodon ''Vulcanodon'' (meaning "volcano tooth") is an extinct genus of sauropod dinosaur from the Early Jurassic Uppermost Forest Sandstone of southern Africa. The only known species is ''V. karibaensis''. Discovered in 1969 in Zimbabwe, it was regard ...
'' and ''
Rhoetosaurus ''Rhoetosaurus'' (meaning "Rhoetos lizard"), named after Rhoetus, a titan in Greek mythology, is a genus of sauropod dinosaur from the Jurassic ( Oxfordian) Walloon Coal Measures of what is now eastern Australia. ''Rhoetosaurus'' is estimated ...
''. The Eusauropoda was coined in 1995 by Paul Upchurch to create a monophyletic new taxonomic group that would include all sauropods, except for the vulcanodontids. Eusauropoda are herbivorous, quadrupedal, and have long necks. They have been found in South America, Europe, North America, Asia, Australia, and Africa. The temporal range of Eusauropoda ranges from the early Jurassic to the Latest Cretaceous periods. The most basal forms of eusauropods are not well known and because the cranial material for the ''Vulcanodon'' is not available, and the distribution of some of these shared derived traits that distinguish Eusauropoda is still completely clear. However the discovery of the vulcanodont,
Tazoudasaurus ''Tazoudasaurus'' is a genus of gravisaurian, probably a vulcanodontidae, vulcanodontid sauropod dinosaurs hailing from the late Early Jurassic (Toarcian), that was recovered in the "Toundoute Continental Series" (Azilal Formation) located in the ...
, which does preserve cranial material did help refine some of the characteristics.


Description

Eusauropods are long-necked, strictly
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
, obligate quadrupedals. They have a highly specialized set of skeletal adaptions due to their large size, and are graviportal.


Teeth and mouth

Yates and Upchurch described eusauropod evolution as moving towards a “bulk-browsing mode of feeding”. They describe the development of lateral plates on the alveolar margins of tooth-bearing bones. These plates can be used to strip foliage, the eusauropod's “U-shaped” jaws create a wide bite, and their loss of “fleshy cheeks” increased the gape. The crowns of eusauropod teeth also have “wrinkled enamel textures”, but it is unclear what this meant for their feeding habits.


Head and neck

The skull length of the basal eusauropod, ''
Patagosaurus ''Patagosaurus'' (meaning "Patagonia lizard") is an extinct genus of eusauropod dinosaur from the Middle-Late Toarcian of Patagonia, Argentina. It was first found in deposits of the Cañadón Asfalto Formation, which date to around 179 to 177&n ...
'' is about . One of the most basal eusauropods, ''
Shunosaurus ''Shunosaurus'', meaning "Lizard from Sichuan", is a genus of sauropod dinosaur from Late Jurassic ( Oxfordian) beds in Sichuan Province in China, from 161 to 157 Million years ago. The name derives from "Shu", an ancient name for the Sichuan pr ...
'', has two characteristic features of the eusauropod elongated neck: the incorporation of the equivalent of the first dorsal vertebra into the cervical region of the spine, and the addition of two cervical vertebra in the middle of the cervical vertebrae. Other
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
of Eusauropoda includes a retracted position of the external nares. Unlike
prosauropods Sauropodomorpha ( ; 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 lo ...
and
theropods Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
, which have a snout with smooth, unprotruding alveolar and subnarial regions, eusauropods have snouts with “stepped anterior margins”. Further distinguishing features of eusauropods include the absence of the contact between the squamosal and the quadratojugal, the absence of the anterior process of the prefrontal, and a distally elongated anterior ramus of the quadratojugal. Separating the anteroventral process of the nasal from the posterolateral process of the premaxilla, eusauropods also have a long maxilla that forms the posteroventral margin of the external naris.


Feet and limbs

Eusauporoda are also hypothesized to have a ''semi-digitigrade'' foot posturem demonstrated by footprint evidence. Paleontologist Jeffrey Wilson explains that eusauropods differ from
theropods Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
and
prosauropods Sauropodomorpha ( ; 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 lo ...
that have digitigrade pes where their heel and metatarsals are lifted off the ground. Eusauropods show asymmetry in their metatarsal shaft diameters where metatarsal I is broader than the others, suggesting that their weight was mostly assumed by their inner feet. According to Steven Salisbury and Jay Nair, basal eusauropods retain four pedal unguals but reduce their phalangeal number in their fourth digit to three units. The metatarsus in eusauropods is less than a quarter of their tibial length, unlike sauropod outgroups that have long hindlimbs and metatarsus that are almost half of their tibial length.


Distribution

Eusauropods are found on all major continents, with diplocodoids being widespread in the
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
, and titanosaurs being found in Southern Hemisphere. However, basal eusauropods that do not fall into either group are fairly well represented. Early eusauropods such as ''Volkeimeria'' and ''Amygdalodon'', and more derived eusauropods such as ''
Patagosaurus ''Patagosaurus'' (meaning "Patagonia lizard") is an extinct genus of eusauropod dinosaur from the Middle-Late Toarcian of Patagonia, Argentina. It was first found in deposits of the Cañadón Asfalto Formation, which date to around 179 to 177&n ...
'' have been found in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. ''Volkeimeria'' is classified as a basal eusauropod, though in 2004 Paul Upchurch was suspicious of its placement, because of its “opisthoceolous cervical centra, the absence of a femoral anterior trochanter, and laterally projecting cnemial crest of the tibia”, and instead thought it may be a generic sauropod. African eusauropods may include ''
Spinophorosaurus ''Spinophorosaurus'' is a genus of sauropod dinosaur that lived in what is now Niger during the Middle Jurassic period. The first two specimens were excavated in the 2000s by German and Spanish teams under difficult conditions. The skeletons were ...
'', from Niger, although that taxon may instead be closer to ''Vulcanodon'' and outside Eusauropoda. Also, ''
Atlasaurus ''Atlasaurus'' is a genus of sauropod dinosaurs from Middle Jurassic (Bathonian to Callovian stages) beds in North Africa.M. Monbaron, D. A. Russell, and P. Taquet. (1999). ''Atlasaurus imelakei'' n.g., n.sp., a brachiosaurid-like sauropod from ...
'' was found in
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
, and ''
Jobaria ''Jobaria'' is a genus of sauropod dinosaur that lived in what is now Niger during the middle Jurassic Period, between 164 and 161 million years ago. ''Jobaria'' is currently the only known valid sauropod from the Tiouraren, where it was disco ...
'' was found in Niger. However, both genera have been found as possible Macronarians, but ''
Atlasaurus ''Atlasaurus'' is a genus of sauropod dinosaurs from Middle Jurassic (Bathonian to Callovian stages) beds in North Africa.M. Monbaron, D. A. Russell, and P. Taquet. (1999). ''Atlasaurus imelakei'' n.g., n.sp., a brachiosaurid-like sauropod from ...
'' was found to be a turiasaurian, and ''Jobaria'' a eusauropod, by a phylogenetic analysis of Xing in 2012. In Europe, the clade
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 ...
has been found in France, Spain, and possibly England, with multiple genera from the same locality in Spain. ''
Cetiosaurus ''Cetiosaurus'' ( meaning 'whale lizard', from the Greek '/ meaning 'sea monster' (later, 'whale') and '/ meaning 'lizard'), is a genus of herbivorous sauropod dinosaur from the Middle Jurassic Period, living about 171 to 165 million years ago ...
'' skeletons have also been found in England, along with the possibly eusauropod genera ''
Cardiodon ''Cardiodon'' (meaning "heart tooth", in reference to the shape) was a herbivorous genus of sauropod dinosaur, based on a tooth from the late Bathonian-age Middle Jurassic Forest Marble Formation of Wiltshire, England. Historically, it is very ...
'' and ''
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 ...
'', known only from teeth. The family
Mamenchisauridae Mamenchisauridae is a family of sauropod dinosaurs belonging to the Eusauropoda known from the Jurassic and Early Cretaceous of Asia and Africa. Mamenchisaurids are characterized by their proportionately extremely long necks. Some members of the ...
is found widespread throughout Asia. A majority of the genera are found in China, although a possible specimen of ''
Mamenchisaurus ''Mamenchisaurus'' ( , or spelling pronunciation ) is an extinct genus of sauropod dinosaurs known for their remarkably long necks which made up nearly half the total body length. Numerous species have been assigned to the genus; however, the val ...
'' has been found in
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
. Also in China, the basal eusauropod ''Nebulasaurus taito'' was found to be a sister taxon to ''Spinophorosaurus'', and more derived than Mamenchisauridae, but less derived than ''Patagosaurus'', and the genus ''
Shunosaurus ''Shunosaurus'', meaning "Lizard from Sichuan", is a genus of sauropod dinosaur from Late Jurassic ( Oxfordian) beds in Sichuan Province in China, from 161 to 157 Million years ago. The name derives from "Shu", an ancient name for the Sichuan pr ...
'' is likely one of the most basal eusauropods. The genus ''
Barapasaurus ''Barapasaurus'' ( ) is a genus of Basal (phylogenetics), basal sauropod dinosaur from Jurassic rocks of India. The only species is ''B. tagorei''. ''Barapasaurus'' comes from the lower part of the Kota Formation, which is of Early to Middle J ...
'' has been found in India, and may represent a cetiosaurid, a basal eusauropod, or a genus outside Eusauropoda.


Paleobiology

The data around sauropods evolution, as Novas points out, is largely based on a few formations mostly in the Northern Hemisphere. However, other beds in places such as Tanzania, specifically the Canadon Asfalto and Canadon Calcereo formations reveal a more diverse and widespread peleobiology of eusauropods in the Late Jurassic period.


Classification

Cladogram of Eusauropoda after Gomez et al. (2024):Cladogram of Sauropoda after Holwerda et al. (2021):Cladogram after Li et al. (2025):


References

{{Taxonbar, from=Q1936183 Sauropoda Dinosaur clades