Ornithosuchidae
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Ornithosuchidae is an extinct
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
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 ...
archosaurs (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 ...
) from the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
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 (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
and Norian stages of the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic 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 European mainland, continental mainland. It comprises England, Scotlan ...
. 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, Dynamosuchus,'' and '' Riojasuchus'' are presently known. The family was first erected by German paleontologist Friedrich von Huene 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
(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 ...
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. 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 Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fis ...
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 separ ...
, 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 a ...
s of the skull is small or absent, excluded by a large contact between the frontals and lacrimals. In other archosaurs, including rauisuchians,
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order (biology), order Aetosauria (; from Ancient Greek, Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized Omnivore, omnivorous or Herbivore, herbivoro ...
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 ...
s, and dinosauromorphs, 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 (neck), 14-15 dorsal (back), 3 sacral (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 characteristi ...
. 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 amphi ...
. 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 wit ...
(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 i ...
(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 sauropod Sauropoda (), whose members are known as sauropods (; ...
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. 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 Suchia, 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