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''Sanjuansaurus'' (" San Juan Province lizard") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of herrerasaurid
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from 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 ...
(
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
) Cancha de Bochas and La Peña Members of the
Ischigualasto Formation The Ischigualasto Formation is a Late Triassic fossiliferous formation and Lagerstätte in the Ischigualasto-Villa Unión Basin of the southwestern La Rioja Province and northeastern San Juan Province in northwestern Argentina. The formation da ...
of the
Ischigualasto-Villa Unión Basin The Ischigualasto-Villa Unión Basin ( es, Cuenca de Ischigualasto-Villa Unión) is a small sedimentary basin located in the Argentine Northwest, Argentina. It is located in the southwestern part of La Rioja Province and the northeastern part of ...
in northwestern Argentina.


Description and classification

''Sanjuansaurus'' was comparable in size to a medium-sized ''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
'', with a
thigh bone The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
that was long and a
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
that is in length (in the description paper the tibia is listed with 260 mm in length due to a typo). Alcober and Martinez performed a
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
and found ''Sanjuansaurus'' to be a herrerasaurid. It was determined that ''Sanjuansaurus'' and''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
'' share many similarities in the morphology of the skull, neck vertebrae, back vertebrae, hip vertebrae, scapula, and the hip bones. Alcober and Martinez observed that ''Sanjuansaurus'' and'' Staurikosaurus ''share many similarities in the morphology of the hip bones, and the
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
. The pubis of ''Sanjuansaurus'', unlike in other herrerasaurids, points toward the cranium.


Distinguishing anatomical features

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group. According to Alcober and Martinez (2010), ''Sanjuansaurus'' can be distinguished based on the following characteristics: * the
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
have shelf-like, posterolaterally directed transverse processes * the neural spines of the sixth to eighth
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
, at least, bear acute anterior and posterior processes * the
glenoid The glenoid fossa of the scapula or the glenoid cavity is a bone part of the shoulder. The word ''glenoid'' is pronounced or (both are common) and is from el, gléne, "socket", reflecting the shoulder joint's ball-and-socket form. It is a sha ...
has everted lateral margins * the pubis is relatively short, measuring 63% of the length of the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
* a pronounced, rugose scar is present on the medial surface of the femur at the level of the fourth
trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...


Discovery

''Sanjuansaurus'' was named and described in
2010 File:2010 Events Collage New.png, From top left, clockwise: The 2010 Chile earthquake was one of the strongest recorded in history; The Eruption of Eyjafjallajökull in Iceland disrupts air travel in Europe; A scene from the opening ceremony of ...
by Oscar Alcober and Ricardo Martinez. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
was named ''S. gordilloi'' after Raul Gordillo, the head fossil preparator and artist of the San Juan Museum. It is known from and based on an associated and partially articulated partial skeleton ( PVSJ 605) consisting of a
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
fragment, most of the
vertebral column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a vertebrate in which the notochord (a flexible rod of uniform composition) found in all chordata, ...
from the
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis * Axis (mathematics), a designator for a Cartesian-coordinat ...
to the
anterior Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
tail, the shoulder blades, an
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
, part of the
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
, most of the
long bone The long bones are those that are longer than they are wide. They are one of five types of bones: long, Short bone, short, Flat bone, flat, Irregular bone, irregular and Sesamoid bone, sesamoid. Long bones, especially the femur and tibia, are subj ...
s of the legs, and a few other bones.


Paleoecology


Provenance and occurrence

''Sanjuansaurus'' holotype specimen PVSJ 605 was discovered in 1994, in gray-green sandstone of the Cancha de Bocas Member near the base of the
Ischigualasto Formation The Ischigualasto Formation is a Late Triassic fossiliferous formation and Lagerstätte in the Ischigualasto-Villa Unión Basin of the southwestern La Rioja Province and northeastern San Juan Province in northwestern Argentina. The formation da ...
, in
Ischigualasto Provincial Park Ischigualasto Provincial Park ( es, Parque Provincial Ischigualasto), also called ''Valle de la Luna'' ("Valley of the Moon" or "Moon Valley"), due to its moon like appearance, is a provincial protected area in the north-east of San Juan Provin ...
in San Juan,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. This horizon dates to approximately 231.4  Ma, during the late
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
Stage of the Late Triassic.


Fauna and habitat

In the Ischigualasto Formation, dinosaurs constituted only about 6% of the total number of fossils, but by the end of the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
Period, dinosaurs were becoming the dominant large land animals, and the other archosaurs and synapsids declined in variety and number. Studies suggest that the
paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
of the Ischigualasto Formation was a volcanically active floodplain covered by forests and subject to strong seasonal rainfalls. The climate was moist and warm, though subject to seasonal variations. Vegetation consisted of
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s (''
Cladophlebis ''Cladophlebis'' is an extinct form genus of fern, used to refer to Paleozoic and Mesozoic fern leaves that have "fern fronds with pinnules that are attached to the rachis, and have a median vein that runs to the apex of the pinnule, and veins ...
''),
horsetail ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Eq ...
s, and giant
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s (''Protojuniperoxylon''). These plants formed
highland Highlands or uplands are areas of high elevation such as a mountainous region, elevated mountainous plateau or high hills. Generally speaking, upland (or uplands) refers to ranges of hills, typically from up to while highland (or highlands) is ...
forests along the banks of rivers. ''Sanjuansaurus'' lived in the jungles of Late Triassic
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
alongside early dinosaurs, ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwest ...
'', ''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
'', ''
Chromogisaurus ''Chromogisaurus'' is an extinct genus of saturnaliid sauropodomorph which existed in Argentina during the Late Triassic (Carnian) period. It is from the Cancha de Bochas, Valle Pintado member of the Ischigualasto Formation. It was a herbivore ...
'', and '' Panphagia'', as well as ''
Saurosuchus ''Saurosuchus'' (meaning "lizard crocodile") is an extinct genus of large loricatan pseudosuchian archosaur that lived in South America during the Late Triassic period. It was a heavy, ground-dwelling, quadrupedal carnivore, likely being the ape ...
'', a giant land-living
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians ...
n (a quadrupedal meat eater with a
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
-like skull); the broadly similar but smaller ''
Venaticosuchus ''Venaticosuchus'' is a genus of pseudosuchian archosaurs from the family Ornithosuchidae. Known from a single species, ''Venaticosuchus rusconii,'' this genus is described based on an incomplete skull and jaw (as well as a lost partial forelimb ...
'', an
ornithosuchid Ornithosuchidae is an extinct family of pseudosuchian archosaurs (distant relatives of modern crocodilians) from the Triassic period. Ornithosuchids were quadrupedal and facultatively bipedal (e.g. like chimpanzees), meaning that they had the a ...
; and the predatory chiniquodontids. Herbivores were much more abundant than carnivores and were represented by
rhynchosaur Rhynchosaurs are a group of extinct herbivorous Triassic archosauromorph reptiles, belonging to the order Rhynchosauria. Members of the group are distinguished by their triangular skulls and elongated, beak like premaxillary bones. Rhynchosaurs ...
s such as ''
Hyperodapedon ''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and namin ...
'' (a beaked reptile);
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order Aetosauria (; from Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized omnivorous or herbivorous pseudosuchians, part of the branch of archosaurs ...
s (spiny armored reptiles);
kannemeyeriid ''Kannemeyeria'' is a genus of dicynodont that lived during the Anisian age of Middle Triassic period in what is now Africa and South America. The generic name is given in honor of Dr. Daniel Rossouw Kannemeyer, the South African fossil collect ...
dicynodonts Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were Herbivore, herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horn ...
(stocky, front-heavy beaked quadrupedal animals) such as ''
Ischigualastia ''Ischigualastia'' is an extinct genus of large dicynodont therapsids that lived during the Late Carnian age and the Early Norian age of the Late Triassic Period. The genus was found in and named after the Ischigualasto Formation (Cancha de Boc ...
''; and
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
traversodontids Traversodontidae is an extinct family of herbivorous cynodonts. Traversodonts were primarily Gondwanan, with many species known from Africa and South America. Recently, traversodonts have also been found from Europe and eastern North America. ...
(somewhat similar in overall form to dicynodonts, but lacking beaks) such as ''
Exaeretodon ''Exaeretodon'' is an extinct genus of fairly large, low-slung traversodontid cynodonts from the southern parts of Pangea. Four species are known, from various formations. ''E. argentinus'' is from the Carnian-age (Late Triassic) Cancha de Boch ...
''. These non-dinosaurian herbivores were much more abundant than early ornithischian dinosaurs like ''
Pisanosaurus ''Pisanosaurus'' () is an extinct genus of early dinosauriform, likely an ornithischian or silesaurid, from the Late Triassic of Argentina. It was a small, lightly built, ground-dwelling herbivore, that could grow up to an estimated long. Onl ...
''.


References

{{Taxonbar, from=Q2073291 Prehistoric saurischians Dinosaur genera Carnian genera Late Triassic dinosaurs of South America Triassic Argentina Fossils of Argentina Ischigualasto Formation Fossil taxa described in 2010