Ornithosuchidae

Last updated

Ornithosuchids
Temporal range: Late Triassic, 228–203.6  Ma
Ornithosuchus1DB.jpg
Life restoration of Ornithosuchus scavenging on the rhynchosaur Hyperodapedon
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Family: Ornithosuchidae
von Huene, 1908
Type species
Ornithosuchus woodwardi
Newton, 1894
Genera

Aenigmaspina ?
Dynamosuchus [1]
Ornithosuchus
Riojasuchus
Venaticosuchus

Contents

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 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 and Norian stages of the Late Triassic with members known from Argentina, Brazil, and the United Kingdom. Four genera, comprising Ornithosuchus , Venaticosuchus, Dynamosuchus, [1] and Riojasuchus are presently known. [2] The family was first erected by German paleontologist Friedrich von Huene in 1908. [3]

Description

Skull

The skull of Riojasuchus Riojasuchus.png
The skull of Riojasuchus

Ornithosuchids can be identified by the presence of an arched diastema, a gap between the teeth at the front of the snout. When the jaw is closed, two large, curved, dentary (lower jaw) teeth fit into the diastema, which is positioned between the premaxilla and maxilla. Two shallow depressions are 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. [4]

Several other features distinguish ornithosuchids from all other early archosaurs. Ornithosuchus and Riojasuchus both possess a small fenestra, or hole, between the palatine and pterygoid bones of the palate, i.e. the roof of the mouth. The contact between the nasal and prefrontal bones of the skull is small or absent, excluded by a large contact between the frontals and lacrimals. In other archosaurs, including rauisuchians, aetosaurs, pterosaurs, and dinosauromorphs, the nasal-prefrontal contact separates the frontal from the lacrimal. [4]

Postcranial skeleton

The postcranial skeleton is nearly completely known in Riojasuchus, incomplete in Ornithosuchus, and entirely unknown in Venaticosuchus. As a result, whether all of the postcranial traits seemingly unique to ornithosuchids actually occurred in all members of the family is uncertain. Ornithosuchids known from decent postcranial remains typically had about 9 cervical (neck), 14-15 dorsal (back), three sacral (hip), and over 20 caudal (tail) vertebrae. Above each vertebra was a pair of bony scutes known as osteoderms. [2]

The femur (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. It was probably an insertion point for m. iliofemoralis cranialis, 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). [5]

Much like the femur, the fibula (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 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 is unclear. [6]

Unlike most other early archosaurs, the pedal unguals (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. [4]

CrocNormal01.png
The "crocodile-normal" ankle of most crurotarsans
CrocReversed01.png
The "crocodile-reversed" ankle of ornithosuchids
   Tibia    Fibula    Astragalus    Calcaneum

Major archosaur groups have often been distinguished from each other based on the structure of their ankles. In most crurotarsans, the astragalus 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 which 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. [4]

Phylogeny

Ornithosuchidae is generally considered to be within the larger clade Suchia, which includes aetosaurs, rauisuchians, and crocodylomorphs. [7] [8] [9] Below is a cladogram based on Nesbitt (2011), showing the placement of Ornithosuchidae in Archosauriformes. [6]

Archosauriformes  

Proterosuchidae ProterosuchusDB flipped.jpg

Erythrosuchus Erythrosuchus.png

Vancleavea Vancleavea white background.jpg

Proterochampsia Pseudochampsa life restoration white background.jpg

Euparkeria Euparkeria white background.png

  Crurotarsi  

Phytosauria Smilosuchus adamanensis flipped.jpg

  Archosauria  

Avemetatarsalia (bird-line archosaurs) Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg

  Pseudosuchia

Ornithosuchidae Ornithosuchus BW white background.jpg

  Suchia  

Gracilisuchus Gracilisuchus BW white background.jpg

Turfanosuchus Turfanosuchus NT White Background.jpg

Revueltosaurus Revueltosaurus.jpg

Aetosauria Desmatosuchus spurensis flipped.jpg

Ticinosuchus Ticinosuchus BW white background.jpg

  Paracrocodylomorpha  

Poposauroidea Poposaurus gracilis (1) flipped.jpg

  Loricata  

Prestosuchus Prestosuchus-chiniquensis (2).jpg

Saurosuchus Saurosuchus BW white background.jpg

Batrachotomus Batrachotomus1DB.jpg

Fasolasuchus

Rauisuchidae Postosuchus kirkpatricki flipped.jpg

Crocodylomorpha Deinosuchus riograndensis.png

Related Research Articles

<span class="mw-page-title-main">Archosaur</span> Group of diapsids broadly classified as reptiles

Archosauria is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being 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 dinosaurs, pterosaurs and extinct relatives of crocodilians. Modern paleontologists define Archosauria as a crown group that includes the most recent common ancestor of living birds and crocodilians, and all of its descendants. The base of Archosauria splits into two clades: Pseudosuchia, which includes crocodilians and their extinct relatives; and Avemetatarsalia, which includes birds and their extinct relatives.

<span class="mw-page-title-main">Crurotarsi</span> Clade of reptiles

Crurotarsi is a clade of archosauriform reptiles that includes crocodilians and stem-crocodilians and possibly bird-line archosaurs too if the extinct, crocodile-like phytosaurs are more distantly related to crocodiles than traditionally thought. Prior to 2011, the group had invariably included only archosaurs closer to crocodilians than to birds and other dinosaurs. An equivalent term for the crocodilian side of the archosaur family tree is Pseudosuchia. This traditional definition of Crurotarsi assumed that phytosaurs were crown-group archosaurs and more closely related to crocodilians than to birds. However, a 2011 study argued that the phytosaur lineage evolved prior to the split between birds and crocodilians. This would mean that phytosaurs were not true archosaurs, and therefore could not be considered representatives of croc-line archosaurs.

<span class="mw-page-title-main">Rauisuchia</span> Informal group of Triassic archosaurs with pillar-erect posture

"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 than to birds and other dinosaurs. First named in the 1940s, Rauisuchia was a name exclusive to Triassic archosaurs which were generally large, carnivorous, and quadrupedal with a pillar-erect hip posture, though exceptions exist for all of these traits. Rauisuchians, as a traditional taxonomic group, were considered distinct from other Triassic archosaur groups such as early dinosaurs, phytosaurs, aetosaurs, and crocodylomorphs.

Phytosaurs are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. and are sometimes referred to as parasuchians. Phytosauria, Parasuchia, Parasuchidae, and Phytosauridae have often been considered equivalent groupings containing the same species. Some recent studies have offered a more nuanced approach, defining Parasuchidae and Phytosauridae as nested clades within Phytosauria as a whole. Phytosaurs were long-snouted and heavily armoured, bearing a remarkable resemblance to modern crocodilians in size, appearance, and lifestyle, as an example of convergence or parallel evolution. The name "phytosaur" means "plant reptile", as the first fossils of phytosaurs were mistakenly thought to belong to plant eaters.

<i>Venaticosuchus</i> Extinct genus of reptiles

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 collected from the Late Triassic (Carnian) Ischigualasto Formation in the Ischigualasto-Villa Unión Basin in northwestern Argentina, which was deposited around 230 million years ago. This fossil material has been termed the holotype specimen PVL 2578. Venaticosuchus incorporated a myriad of features present in the other two genera of ornithosuchids, Ornithosuchus and Riojasuchus. However, it also had several unique traits, relating to the lower jaw.

<i>Gracilisuchus</i> Genus of fossil reptiles

Gracilisuchus is an extinct genus of tiny pseudosuchian from the Late Triassic of Argentina. It contains a single species, G. stipanicicorum, which is placed in the clade Suchia, close to the ancestry of crocodylomorphs. Both the genus and the species were first described by Alfred Romer in 1972.

<i>Riojasuchus</i> Extinct genus of reptiles

Riojasuchus is an extinct genus of ornithosuchid archosaur from the Late Triassic (Norian) of Argentina. Ornithosuchidae was a widespread family of facultatively bipedal pseudosuchians with adaptations for scavenging. Riojasuchus is notable as one of the youngest and most complete members of the family. The type and only known species, Riojasuchus tenuisceps, was named and described by José Bonaparte in 1967. It was one of the first of many well-preserved Triassic archosaurs to be discovered in Argentina. The holotype specimen, PVL 3827, was found in the Los Colorados Formation of the Ischigualasto-Villa Unión Basin in northwestern Argentina.

<span class="mw-page-title-main">Prestosuchidae</span> Extinct family of reptiles

Prestosuchidae is a polyphyletic grouping of carnivorous archosaurs that lived during the Triassic. They were large active terrestrial apex predators, ranging from around 2.5 to 7 metres in length. They succeeded the Erythrosuchidae as the largest archosaurs of their time. While resembling erythrosuchids in size and some features of the skull and skeleton, they were more advanced in their erect posture and crocodile-like ankle, indicating more efficient gait. "Prestosuchids" flourished throughout the whole of the middle, and the early part of the late Triassic, and fossils are so far known from Europe, India, Africa (Tanzania), Argentina, and Paleorrota in Brazil. However, for a long time experts disagree regarding the phylogenetic relationships of the group, what genera should be included, and whether indeed the "Prestosuchidae" constitute a distinct family.

<i>Teleocrater</i> Extinct genus of reptiles

Teleocrater is a genus of avemetatarsalian archosaur from the Middle Triassic Manda Formation of Tanzania. The name was coined by English paleontologist Alan Charig in his 1956 doctoral dissertation, but was only formally published in 2017 by Sterling Nesbitt and colleagues. The genus contains the type and only species T. rhadinus. Uncertainty over the affinities of Teleocrater have persisted since Charig's initial publication; they were not resolved until Nesbitt et al. performed a phylogenetic analysis. They found that Teleocrater is most closely related to the similarly enigmatic Yarasuchus, Dongusuchus, and Spondylosoma in a group that was named the Aphanosauria. Aphanosauria was found to be the sister group of the Ornithodira, the group containing dinosaurs and pterosaurs.

<i>Turfanosuchus</i> Extinct genus of reptiles

Turfanosuchus is a genus of archosauriform reptile, likely a gracilisuchid archosaur, which lived during the Middle Triassic (Anisian) of northwestern China. The type species, T. dabanensis, was described by C.C. Young in 1973, based on a partially complete but disarticulated fossil skeleton found in the Kelamayi Formation of the Turfan Basin.

Yarasuchus is an extinct genus of avemetatarsalian archosaur that lived during the Anisian stage of the Middle Triassic of India. The genus was named and described in 2005 from a collection of disarticulated but fairly complete fossil material found from the Middle Triassic Yerrapalli Formation. The material is thought to be from two individuals, possibly three, with one being much more complete and articulated than the other. The type and only species is Y. deccanensis. Yarasuchus was a quadruped roughly 2–2.5 metres (6.6–8.2 ft) long, with an elongated neck and tall spines on its vertebrae. Unlike other quadrupedal Triassic reptiles, the limbs and shoulders of Yarasuchus were slender, and more like those of ornithodirans.

<i>Mandasuchus</i> Extinct genus of reptiles

Mandasuchus is an extinct genus of loricatan pseudosuchian from the Manda Formation of Tanzania, which dates back to the Anisian stage of the Middle Triassic. Although this genus was first mentioned by Alan Charig in 1956, a formal description was not published until 2018.

<i>Vancleavea</i> Extinct genus of reptiles

Vancleavea is a genus of extinct, armoured, non-archosaurian archosauriforms from the Late Triassic of western North America. The type and only known species is V. campi, named by Robert Long & Phillip A Murry in 1995. At that time, the genus was only known from fragmentary bones including osteoderms and vertebrae. However, since then many more fossils have been found, including a pair of nearly complete skeletons discovered in 2002. These finds have shown that members of the genus were bizarre semiaquatic reptiles. Vancleavea individuals had short snouts with large, fang-like teeth, and long bodies with small limbs. They were completely covered with bony plates known as osteoderms, which came in several different varieties distributed around the body. Phylogenetic analyses by professional paleontologists have shown that Vancleavea was an archosauriform, part of the lineage of reptiles that would lead to archosaurs such as dinosaurs and crocodilians. Vancleavea lacks certain traits which are present in most other archosauriforms, most notably the antorbital, mandibular and supratemporal fenestrae, which are weight-saving holes in the skulls of other taxa. However, other features clearly support its archosauriform identity, including a lack of intercentra, the presence of osteoderms, an ossified laterosphenoid, and several adaptations of the femur and ankle bones. In 2016, a new genus of archosauriform, Litorosuchus, was described. This genus resembled both Vancleavea and more typical archosauriforms in different respects, allowing Litorosuchus to act as a transitional fossil linking Vancleavea to less aberrant archosauriforms.

<span class="mw-page-title-main">Suchia</span> Clade of reptiles

Suchia is a clade of archosaurs containing the majority of pseudosuchians. It was defined as the least inclusive clade containing Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, and Crocodylus niloticus by Nesbitt (2011). Generally the only pseudosuchian group which is omitted from Suchia is the family Ornithosuchidae, although at least one analysis classifies ornithosuchids as close relatives of erpetosuchids and aetosaurs. Phytosaurs are also excluded from Suchia, although it is not certain whether they qualify as pseudosuchians in the first place.

<span class="mw-page-title-main">Loricata</span> Extinct clade of reptiles

Loricata is a clade of archosaur reptiles that includes crocodilians and some of their Triassic relatives, such as Postosuchus and Prestosuchus. More specifically, Loricata includes Crocodylomorpha and most "rauisuchians", a paraphyletic grade of large terrestrial pseudosuchians which were alive in the Triassic period and ancestral to crocodylomorphs.

<span class="mw-page-title-main">Paracrocodylomorpha</span> Clade of reptiles

Paracrocodylomorpha is a clade of pseudosuchian archosaurs. The clade includes the diverse and unusual group Poposauroidea as well as the generally carnivorous and quadrupedal members of Loricata, including modern crocodylians. Paracrocodylomorpha was named by paleontologist J. Michael Parrish in 1993, although the group is now considered to encompass more reptiles than his original definition intended. The most recent definition of Paracrocodylomorpha, as defined by Sterling Nesbitt in 2011, is "the least inclusive clade containing Poposaurus and Crocodylus niloticus. Most groups of paracrocodylomorphs became extinct at the end of the Triassic period, with the exception of the crocodylomorphs, from which crocodylians such as crocodiles and alligators evolved in the latter part of the Mesozoic.

<span class="mw-page-title-main">Erpetosuchidae</span> Extinct family of reptiles

Erpetosuchidae is an extinct family of pseudosuchian archosaurs. Erpetosuchidae was named by D. M. S. Watson in 1917 to include Erpetosuchus. It includes the type species Erpetosuchus granti from the Late Triassic of Scotland, Erpetosuchus sp. from the Late Triassic of eastern United States and Parringtonia gracilis from the middle Middle Triassic of Tanzania; the group might also include Dyoplax arenaceus from the Late Triassic of Germany, Archeopelta arborensis and Pagosvenator candelariensis from Brazil and Tarjadia ruthae from Argentina.

Nundasuchus is an extinct genus of crurotarsan, possibly a suchian archosaur related to Paracrocodylomorpha. Remains of this genus are known from the Middle Triassic Manda beds of southwestern Tanzania. It contains a single species, Nundasuchus songeaensis, known from a single partially complete skeleton, including vertebrae, limb elements, osteoderms, and skull fragments.

Aphanosauria is an extinct group of reptiles distantly related to dinosaurs. They are at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchia, includes modern crocodilians. Aphanosaurs possessed features from both groups, indicating that they are the oldest and most primitive known clade of avemetatarsalians, at least in terms of their position on the archosaur family tree. Other avemetatarsalians include the flying pterosaurs, small bipedal lagerpetids, herbivorous silesaurids, and the incredibly diverse dinosaurs, which survive to the present day in the form of birds. Aphanosauria is formally defined as the most inclusive clade containing Teleocrater rhadinus and Yarasuchus deccanensis but not Passer domesticus or Crocodylus niloticus. This group was first recognized during the description of Teleocrater. Although only known by a few genera, Aphanosaurs had a widespread distribution across Pangaea in the Middle Triassic. They were fairly slow quadrupedal long-necked carnivores, a biology more similar to basal archosaurs than to advanced avemetatarsalians such as pterosaurs, lagerpetids, and early dinosaurs. In addition, they seemingly possess 'crocodile-normal' ankles, showing that 'advanced mesotarsal' ankles were not basal to the whole clade of Avemetatarsalia. Nevertheless, they possessed elevated growth rates compared to their contemporaries, indicating that they grew quickly, more like birds than other modern reptiles. Despite superficially resembling lizards, the closest modern relatives of aphanosaurs are birds.

<i>Dynamosuchus</i> Extinct genus of reptiles

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. Dynamosuchus is considered a close relative of Venaticosuchus, which is known from the Ischigualasto Formation of Argentina. Ornithosuchids are one of many groups which lived in the Santa Maria and Ischigualasto Formations, which formed at approximately the same time and were ecologically similar. As a large scavenging reptile, Dynamosuchus helps to illuminate the trophic structure of the Santa Maria Formation. It also supports the hypothesis that ornithosuchids had diversified throughout South America by the start of the Carnian, and were not originally endemic to the Ischigualasto-Villa Unión Basin.

References

  1. 1 2 Müller, Rodrigo T.; Von Baczko, M. Belén; Desojo, Julia B.; Nesbitt, Sterling J. (31 January 2020). "The first ornithosuchid from Brazil and its macroevolutionary and phylogenetic implications for Late Triassic faunas in Gondwana" (PDF). Acta Palaeontologica Polonica. 65. doi: 10.4202/app.00652.2019 .
  2. 1 2 Baczko, M. Belén von; Ezcurra, Martín D. (2013-01-01). "Ornithosuchidae: a group of Triassic archosaurs with a unique ankle joint". Geological Society, London, Special Publications. 379 (1): 187–202. doi:10.1144/SP379.4. hdl: 11336/41617 . ISSN   0305-8719. S2CID   130687362.
  3. Huene, F. von. 1908. "Die Dinosaurier der europäischen Triasformation mit Berücksichtigung der aussereuropäischen Vorkommnisse". Geologische und Paläontologische Abhandlungen1(Suppl.): 1–419
  4. 1 2 3 4 Sereno, P.C. (1991). "Basal archosaurs: phylogenetic relationships and functional implications". Journal of Vertebrate Paleontology. 11 (Suppl. 4): 1–53. doi:10.1080/02724634.1991.10011426.
  5. Langer, Max C.; Benton, Michael J. (6 November 2006). "Early dinosaurs: A phylogenetic study" (PDF). Journal of Systematic Palaeontology. 4 (4): 309–358. doi:10.1017/s1477201906001970. ISSN   1477-2019. S2CID   55723635.
  6. 1 2 Nesbitt, S.J. (2011). "The early evolution of archosaurs: relationships and the origin of major clades". Bulletin of the American Museum of Natural History. 352: 1–292. doi: 10.1206/352.1 . hdl:2246/6112.
  7. Nesbitt, S.J.; Norrell, M.A. (2006). "Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda)". Proceedings of the Royal Society B. 273 (1590): 1045–1048. doi:10.1098/rspb.2005.3426. PMC   1560254 . PMID   16600879.
  8. Nesbitt, S.J. (2007). "The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa" (PDF). Bulletin of the American Museum of Natural History. 302: 1–84. doi:10.1206/0003-0090(2007)302[1:TAOEOA]2.0.CO;2. hdl:2246/5840. S2CID   55677195.
  9. Brusatte, S.L.; Benton, M.J.; Desojo, J.B.; Langer, M.C. (2010). "The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida)" (PDF). Journal of Systematic Palaeontology. 8 (1): 3–47. doi:10.1080/14772010903537732. hdl: 20.500.11820/24322ff3-e80e-45f2-8d53-d35fd104195c . S2CID   59148006.