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Ornithosuchidae is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
pseudosuchian 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 ...
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 (distant relatives of modern
crocodilians 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 re ...
) from 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. Ornithosuchids were
quadrupedal Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
and facultatively
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
(e.g. like chimpanzees), meaning that they had the ability to walk on two legs for short periods of time. They had distinctive, downturned snouts, unique, "crocodile-reversed" ankle bones, and several other features that distinguish them from other archosaurs. Ornithosuchids were geographically widespread during the
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 ...
and
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
stages 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 ...
with members known from
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 ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
. Four genera, comprising ''
Ornithosuchus ''Ornithosuchus'' (meaning "bird crocodile") is an extinct genus of pseudosuchians from the Late Triassic (Carnian) Lossiemouth Sandstone of Scotland. It was originally thought to be the ancestor to the carnosaurian dinosaurs (such as ''Allosau ...
'', ''
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 ...
,
Dynamosuchus ''Dynamosuchus'' is an extinct genus of pseudosuchian archosaurs from the family Ornithosuchidae. It is known from a single species, ''Dynamosuchus collisensis'', which is based on a partial skeleton from the Santa Maria Formation of Brazil. ''D ...
,'' and ''
Riojasuchus ''Riojasuchus'' is an extinct genus of Late Triassic ( Norian) quadrupedal archosaur. ''Riojasuchus'' is a member of Ornithosuchidae, a family of facultatively bipedal carnivores that were geographically widespread during the Late Triassic. Tw ...
'' are presently known. The family was first erected by German paleontologist
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
in 1908.


Description


Skull

Ornithosuchids can be identified by the presence of an arched
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
, a gap between the teeth at the front of the snout. When the jaw is closed, two large curved
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
(lower jaw) teeth fit into the diastema, which is positioned between the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
and
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
. There are two shallow depressions on the wall of the diastema to accommodate these teeth. The large dentary teeth of ''Ornithosuchus'' and ''Riojasuchus'' are placed behind a smaller procumbent dentary tooth that sticks out from the jaw. This type of tooth position is not seen in any other basal archosaurs. Another characteristic feature of ornithosuchids is their unusual downturned, overhanging snout, seen in ''Riojasuchus'' and ''Venaticosuchus'', but not ''Ornithosuchus''. Several other features distinguish ornithosuchids from all other early archosaurs. ''Ornithosuchus'' and ''Riojasuchus'' both possess a small fenestra, or hole, between the
palatine A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
and
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates, behind the palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the th ...
s of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
, i.e. the roof of the mouth. The contact between the nasal and
prefrontal bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most modern and ...
s of the skull is small or absent, excluded by a large contact between the frontals and
lacrimals The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the Orbit (anatomy), orbit. It has two surfaces and four borders. Several ...
. In other archosaurs, including
rauisuchian "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 ...
s,
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,
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s, and
dinosauromorph Dinosauromorpha is a clade of avemetatarsalian archosaurs (reptiles closer to birds than to crocodilians) that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lag ...
s, the nasal-prefrontal contact separates the frontal from the lacrimal.


Postcranial skeleton

The postcranial skeleton is nearly completely known in ''Riojasuchus'', incomplete in ''Ornithosuchus'', and entirely unknown in ''Venaticosuchus''. As a result, it is uncertain whether all of the postcranial traits seemingly unique to ornithosuchids actually occurred in all members of the family. Ornithosuchids known from decent postcranial remains typically had about 9
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 **cerv ...
(neck), 14-15 dorsal (back), 3
sacral Sacral may refer to: *Sacred Sacred describes something that is dedicated or set apart for the service or worship of a deity; is considered worthy of spiritual respect or devotion; or inspires awe or reverence among believers. The property ...
(hip), and over 20 caudal (tail)
vertebrae 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 i ...
. Above each vertebra was a pair of bony scutes known as
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...
. 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 ...
(thigh bone) has a pronounced anterior trochanter. The anterior trochanter, sometimes known as the "lesser trochanter" (but unrelated to the lesser trochanter of the femur in humans), is a ridge on the outer surface of the femur, near the
femoral head The femoral head (femur head or head of the femur) is the highest part of the thigh bone (femur). It is supported by the femoral neck. Structure The head is globular and forms rather more than a hemisphere, is directed upward, medialward, and a l ...
. It was probably an insertion point for the ''iliofemoralis cranialis'' muscle which helps to raise the leg. Most archosaurs and archosaur relatives lack a distinct anterior trochanter, but ornithosuchids are an exception, along with most dinosauromorphs (dinosaurs and their close relatives). Much like the femur, the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...
(outer shin bone) also has a distinctive point for muscle insertion. The muscle in question is the ''iliofibularis'', which helps to straighten the limbs. In most archosaurs, the ''iliofibularis'' is inserted onto the fibula by means of a tiny ridge on the proximal part of the fibula, near the knee. However, ornithosuchids have a much larger, knob-shaped ''iliofibularis'' insertion point located about midway down the shaft of the fibula. Phytosaurs and aetosaurs also share a knob-like attachment point midway down the fibula, so it is unclear whether the case in ornithosuchids is a unique case of convergent evolution, or alternatively the retention of a trait independently lost by several archosaur lineages. Unlike most other early archosaurs, the pedal
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...
s (the distalmost bones of the feet that form claws) are laterally compressed. They are sharp and recurved. The unguals are very deep, being taller than they are long, especially on the inner digits. This type of claw is not seen in any other Triassic archosaur except for pterosaurs. Major archosaur groups have often been distinguished from each other based on the structure of their ankles. In most crurotarsans, the
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
has a convex projection that fits into a concave space in the
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. St ...
. This condition is often referred to as a "crocodile-normal" ankle, as it is the most common ankle type in crurotarsans. Ornithosuchids are unique among crurotarsans, and all other archosaurs, in their possession of a "crocodile-reversed" ankle. In a "crocodile-reversed" ankle, the placement of the concavity is reversed: instead of being on the calcaneum, it is on the astragalus. In ornithosuchids, the calcaneum bears a convex projection that is analogous to the convex projection on the "crocodile-normal" astragalus.


Phylogeny

Ornithosuchidae is generally considered to be within the larger
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Suchia Suchia is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing '' Aetosaurus ferratus'', '' Rauisuchus tiradentes'', '' Prestosuchus ch ...
, which includes aetosaurs, rauisuchians, and crocodylomorphs. Below is a cladogram based on Nesbitt (2011), showing the placement of Ornithosuchidae in Archosauriformes.


References


External links


Palaeos Vertebrates
{{Taxonbar, from=Q134276 Late Triassic pseudosuchians Late Triassic first appearances Rhaetian extinctions Prehistoric reptile families