Sillosuchus
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''Sillosuchus'' is a genus of
shuvosaurid Shuvosauridae is an extinct family (biology), family of theropod-like pseudosuchians within the clade Poposauroidea. Shuvosaurids existed in North America (United States) and South America (Argentina) during the Late Triassic period (late Carnian ...
poposauroid Poposauroidea is a clade of advanced pseudosuchians (archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as ''Qianosuchus'' and ''Lotosaurus''. Howeve ...
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
that lived in
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 ...
during 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 ...
period. Shuvosaurids were an unusual family of reptiles belonging to the group
Poposauroidea Poposauroidea is a clade of advanced pseudosuchians (archosaurs closer to crocodilians than to dinosaurs). It includes poposaurids, shuvosaurids, ctenosauriscids, and other unusual pseudosuchians such as ''Qianosuchus'' and ''Lotosaurus''. Howeve ...
; although their closest modern relatives are
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
ns, they were bipedal and lightly armored, with
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 ...
-like hip and skull structures. Based on skull remains from members of the family such as ''
Effigia ''Effigia'' was an extinct genus of shuvosaurid known from the Late Triassic of New Mexico, south-western USA. With a bipedal stance, long neck, and a toothless beaked skull, ''Effigia'' and other shuvosaurids bore a resemblance to the ornithomim ...
'', they were also toothless and likely beaked herbivores.


Discovery and naming

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
specimen of ''Sillosuchus'', PVSJ 85, is a partial skeleton discovered in sediments 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 ...
, Cancha de Bochas Member in 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 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 ...
. The skeleton includes various
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e,
ribs The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a semi- ...
, and
pelvic 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 ...
(hip) bones. A
shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of t ...
,
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
(upper arm bone), and partial
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 ...
e (inner shin bones) were also associated with the skeleton but not featured in the original description of the specimen. The generic name refers to Dr. William Sill, and ''suchus'', from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
word meaning "
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
". The specific name, ''longicervix'', refers to the elongated cervical vertebrae of the genus. Later studies have shown that, while the neck of ''Sillosuchus'' was long compared to most pseudosuchians, this trait was par for the course for shuvosaurids. A reconstruction of the fossil was presented in 2008 by the
National University of San Juan The National University of San Juan (in Castilian, ''Universidad Nacional de San Juan'', UNSJ) is a public university in Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the ...
in
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 ...
, in an homage to Dr. William Dudley Sill who was an important promoter of paleontology in
San Juan San Juan, Spanish for Saint John, may refer to: Places Argentina * San Juan Province, Argentina * San Juan, Argentina, the capital of that province * San Juan, Salta, a village in Iruya, Salta Province * San Juan (Buenos Aires Underground), ...
. ''Sillosuchus'' is the only named genus of bipedal poposauroid currently known from outside
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. Other bipedal poposauroids included the related shuvosaurids ''
Shuvosaurus ''Shuvosaurus'' (meaning "Shuvo's lizard") is a genus of beaked reptile from the Late Triassic of western Texas. Despite looking superficially similar to a theropod dinosaur, it is actually more closely related to crocodilians. Discovery and ...
'' and ''Effigia,'' as well as the carnivorous ''
Poposaurus ''Poposaurus'' (" Popo Agie reptile") is an extinct genus of pseudosuchian archosaur from the Late Triassic of the southwestern United States. It belongs to the clade Poposauroidea, an unusual group of Triassic pseudosuchians that includes sail-b ...
''.


Description

The individual of ''Sillosuchus'' to which the original specimen belonged had an estimated length of about . However, other remains indicate that it could grow much larger. Most notable among additional remains was PVL 2472, an isolated
cervical In anatomy, cervical is an adjective that has two meanings: # of or pertaining to any neck. # of or pertaining to the female cervix: i.e., the ''neck'' of the uterus. *Commonly used medical phrases involving the neck are **cervical collar **cervic ...
(neck vertebra) initially misidentified as belonging to the giant quadrupedal predator ''
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 distant relative of ''Sillosuchus''. This bone was long, while cervicals from the holotype were only long. Using its more complete relatives to estimate length, this bone likely belonged to a ''Sillosuchus'' that was in length. This would indicate that ''Sillosuchus'' was among the largest terrestrial
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
ns (crocodilian-line
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
s). ''Sillosuchus'' had several unique features compared to its relatives. The neck and back vertebrae had large excavations or pockets on the side, an unusual trait that assisted paleontologists in assigning the giant cervical vertebra to ''Sillosuchus'' rather than ''Saurosuchus''. The left and right
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to west ...
(rear-pointing hip bones) were short and fused to each other. They were also flattened in a dorsoventral (top-to-bottom) direction, unlike the case in most other reptiles in which they were flattened in a mediolateral (side-to-side) direction. Otherwise, the hip is practically identical to that of ''Effigia'' and ''
Shuvosaurus ''Shuvosaurus'' (meaning "Shuvo's lizard") is a genus of beaked reptile from the Late Triassic of western Texas. Despite looking superficially similar to a theropod dinosaur, it is actually more closely related to crocodilians. Discovery and ...
'' (also sometimes known as "''Chatterjeea''").


References

{{Portal bar, Paleontology Poposauroids Carnian genera Late Triassic reptiles of South America Triassic Argentina Fossils of Argentina Ischigualasto Formation Fossil taxa described in 1997 Prehistoric pseudosuchian genera