Antetonitrus
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''Antetonitrus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of sauropodiform
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 ...
found in the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassicâ ...
Elliot Formation The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern Eastern Cape, so ...
of
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. The only species is ''Antetonitrus ingenipes''. Sometimes considered a basal
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 ...
, it is crucial for the understanding of the origin and early evolution of this group. It was a
quadruped Quadrupedalism is a form of locomotion in which animals have four legs that are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four legs is said to be a quadruped (fr ...
al
herbivore 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 ...
, like its later relatives, but shows primitive adaptations to use the forelimbs for grasping, instead of purely for weight support.


Discovery and naming

Adam Yates, an
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
n expert on early
sauropodomorph 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 ...
s, named ''Antetonitrus'' in a 2003 report co-authored by
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
n James Kitching. The name is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''ante-'' ("before") and ''tonitrus'' ("thunder"), which refers to its existence, before other known sauropods, specifically ''
Brontosaurus ''Brontosaurus'' (; meaning "thunder lizard" from the Ancient Greek, Greek words , "thunder" and , "lizard") is a genus of herbivorous sauropod dinosaur that lived in present-day United States during the Late Jurassic period. It was describe ...
'' ("thunder lizard"). The one known species of ''Antetonitrus'' is called ''A. ingenipes'', from the Latin ''ingens'' ("massive") and ''pes'' ("foot"), because it shows the beginning of the development of feet designed solely to support weight. The
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s now known as ''Antetonitrus'' were discovered by Kitching in 1981 in the Free State of South Africa, and were stored in the Bernard Price Institute where they were labeled as '' Euskelosaurus''. Yates recognized them as a separate taxon and published a description several years later. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, or original specimen, consists of several
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e and numerous bones from both forelimb and hind limb, all presumed to be from one individual. Five more limb bones from another smaller individual were also referred to the genus.


Description

The holotype specimen may have measured in length, in hip height and in body mass. However, the
neural arch Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
es of the vertebrae were not fused with the centra, indicating that this individual perhaps was not fully grown. ''Antetonitrus'' shows several features which appear to be similar to those of later sauropods, but still retains some primitive features. Unlike most of its smaller and more lightly built ancestors, ''Antetonitrus'' was primarily quadrupedal. Like sauropods, its forelimbs were much longer relative to its hind legs than earlier animals, while the
metatarsus The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
was shortened. However, the first digit of the hand, also called the "thumb" or pollex, was still twisted and flexible, capable of grasping against the hand. In more derived sauropods, the wrist bones are large and thick, arranged in such a way as to lock the hand into a permanently pronated position for full-time weight support, and the hand is incapable of grasping. ''Antetonitrus'' already shows adaptations for an increasing body size as seen in all later sauropods: The wrist bones were broader and thicker to support more weight, whereas the femur was elliptical in cross section. The vertebrae bear high neural spines and well developed hyposphene-hypantrum articulations which add rigidity to the trunk. The first toe of the hind foot already bears a large claw longer than the first metatarsal; however, this claw was not sickle shaped as seen in later sauropods. The femur was slightly sigmoidal (S-curved) in lateral view rather than straight as in other sauropods. Despite being a true sauropod, the long bone development of ''Antetonitrus'' was more characteristic of basal sauropodomorphs, with subadults possessing, as derived sauropods do, highly vascularised fibrolamellar bone throughout their cortices, but with intermittent growth lines as seen in more basal sauropodomorphs instead of the absence of growth lines or presence of growth lines exclusively in the outer circumferential layer.


Classification

A
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis by Yates and Kitching recognizes ''Antetonitrus'' as a basal sauropod, occupying a position between more derived animals such as ''
Isanosaurus ''Isanosaurus'' ("North-eastern Thailand lizard") is an extinct genus of sauropod dinosaur from Thailand. It was originally dated to approximately 219-199  Mya (unit), million years ago during the Late Triassic (late Norian to Rhaetian stage ...
'' or ''
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 more basal sauropods like '' Melanorosaurus''. The back vertebrae are extremely similar to '' Lessemsaurus'' from South America, while the limb bones are similar to ''
Blikanasaurus ''Blikanasaurus'' is a genus of sauropodomorph dinosaur from the late Triassic of South Africa. The generic name ''Blikanasaurus'' is derived from Greek, meaning "lizard from Blikana". The species name ''cromptoni'' is taken from the surname of ...
'', another stocky early sauropod from South Africa. However, these animals were not included in a cladistic analysis with ''Antetonitrus'' because they are poorly known. Apaldetti et al. (2018) erected Lessemsauridae, a
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 ...
containing ''Antetonitrus'', '' Lessemsaurus'' and '' Ingentia''. Their cladogram is reproduced below: The following cladogram shows the position of ''Antetonitrus'' within Massopoda, according to Oliver W. M. Rauhut and colleagues, 2020:


Paleogeography

While ''Antetonitrus'' is not the earliest sauropod from a
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
standpoint, it is currently one of the oldest known sauropod
chronologically Chronology (from Latin , from Ancient Greek , , ; and , ''-logia'') is the science of arranging events in their order of occurrence in time. Consider, for example, the use of a timeline or sequence of events. It is also "the determination of t ...
, or rather tied for that distinction with other early sauropods from the same formation, like ''Melanorosaurus'' and ''Blikanasaurus''. Fossils of these animals were recovered from the
Elliot Formation The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern Eastern Cape, so ...
. Initially it was thought to have been recovered from the
Lower Elliot Formation The Elliot Formation is a geological formation and forms part of the Stormberg Group, the uppermost geological Stratigraphic unit, group that comprises the greater Karoo Supergroup. Outcrops of the Elliot Formation have been found in the northern ...
which dates to the
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
stage of the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
, or approximately 221 to 210
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
. Later studies indicate that it was actually recovered from the Early Jurassic Upper Elliot Formation. Before ''Antetonitrus'' and the other animals recovered from the Elliot Formation were recognized as sauropods, the oldest known sauropod had been ''Isanosaurus'' from the
Rhaetian The Rhaetian is the latest age (geology), age of the Triassic period (geology), Period (in geochronology) or the uppermost stage (stratigraphy), stage of the Triassic system (stratigraphy), System (in chronostratigraphy). It was preceded by the N ...
stage of
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 ...
. Early sauropods and their
sauropodomorph 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 ...
relatives were found around the world as all of the continents were at the time united into the single supercontinent,
Pangaea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous period approximately 335 mi ...
, which made dispersal across the entire terrestrial world possible.


References


External links


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, written by Adam Yates {{Taxonbar, from=Q772043 Sauropoda Dinosaur genera Hettangian dinosaurs Elliot Formation Dinosaurs of South Africa Fossil taxa described in 2003 Taxa named by James Kitching