Crocodylomorphs Temporal range: Late Triassic –Present, | |
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Skeleton of Terrestrisuchus , an early saltoposuchid crocodylomorph | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Paracrocodylomorpha |
Clade: | Loricata |
Clade: | Crocodylomorpha Hay, 1930 |
Subgroups | |
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Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. Extinct crocodylomorphs were considerably more ecologically diverse than modern crocodillians. The earliest and most primitive crocodylomorphs are represented by "sphenosuchians", a paraphyletic assemblage containing small-bodied, slender forms with elongated limbs that walked upright, which represents the ancestral morphology of Crocodylomorpha. These forms persisted until the end of the Jurassic. [4] During the Jurassic, crocodylomorphs morphologically diversified into numerous niches, with the subgroups Neosuchia (which includes modern crocodilians) and the extinct Thalattosuchia adapting to aquatic life, while some terrestrial groups adopted herbivorous and omnivorous lifestyles. Terrestrial crocodylomorphs would continue to co-exist alongside aquatic forms until becoming extinct during the Miocene.
The earliest lineages of Crocodylomorpha are placed into the paraphyletic "Sphenosuchia", which are charactersed by slender bodies with elongate legs. The oldest known crocodylmorph is Trialestes , known from the Late Triassic (Carnian-Norian) of Argentina, around 231–225 million years ago, [5] the last groups of "sphenosuchians" persisted until the end of the Jurassic. [4] During the Jurassic, crocodylomorphs diversified, including the emergence of herbivorous and omnivorous forms, [6] as well as the aquatically adapted Neosuchia and Thalattosuchia, with Thalattosuchia and several groups of neosuchians becoming adapted to a marine lifestyle over the Jurassic and Cretaceous [7] During the Cretaceous, the Notosuchia were a diverse group across the Southern Hemisphere occupying many diverse ecologies. [8] Modern crocodilians, a subgroup of Neosuchia, emerged during the Late Cretaceous. [7] Crocodylomorph diversity was severely reduced by the end-Cretaceous extinction event. [8] The last group of terrestrially adapted crocodylomorphs was the Sebecidae, a group of large predatory notosuchians which persisted in South America until the middle Miocene around 12 million years ago. [9] [7]
Historically, all known living and extinct crocodiles were indiscriminately lumped into the order Crocodilia. However, beginning in the late 1980s, many scientists began restricting the order Crocodilia to the living species and close extinct relatives such as Mekosuchus . The various other groups that had previously been known as Crocodilia were moved to Crocodylomorpha and the slightly more restrictive Crocodyliformes. [10] Crocodylomorpha has been given the rank of superorder in some 20th and 21st century studies. [11]
The old Crocodilia was subdivided into the suborders:
Mesosuchia is a paraphyletic group as it does not include eusuchians (which nest within Mesosuchia). Mesoeucrocodylia was the name given to the clade that contains mesosuchians and eusuchians (Whetstone and Whybrow, 1983).
Below is a cladogram modified from Nesbitt (2011) [12] and Bronzati (2012). [13]
Crocodylomorpha | |
The previous definitions of Crocodilia and Eusuchia did not accurately convey evolutionary relationships within the group. The only order-level taxon that is currently considered valid is Crocodilia in its present definition. Prehistoric crocodiles are represented by many taxa, but since few major groups of the ancient forms are distinguishable, a conclusion on how to define new order-level clades is not yet possible. (Benson & Clark, 1988).
The Crocodylomorpha comprise a variety of forms, shapes, and sizes, which occupied a range of habitats. As with most amniotes, Crocodylomorphs were and are oviparous, laying eggs in a nest or mound, known from strata as old as the Late Jurassic. [14] Adult size varies widely, from about 55 cm long in Knoetschkesuchus to much larger dimensions, as in Sarcosuchus . Most crocodylomorphs were carnivores, but many lineages evolved to be obligate piscivores, such as the extant gharials.
In some forms, like Hesperosuchus and Terrestrisuchus, metatarsal V still had one or two phalanges, [15] but in Crocodyliformes all metatarsal V phalanges have been lost. [16]
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.
"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.
Saltoposuchus is an extinct genus of small, long-tailed crocodylomorph reptile (Sphenosuchia), from the Norian of Europe. The name translated means "leaping foot crocodile". It has been proposed that Terrestrisuchus gracilis and Saltoposuchus connectens represent different ontogenetic stages of the same genus. Saltoposuchus was commonly referred to in popular literature as the ancestor to dinosaurs; however, recent scientific research shows that this is not the case.
Goniopholis is an extinct genus of goniopholidid crocodyliform that lived in Europe and North America during the Late Jurassic and Early Cretaceous. Like other goniopholidids, it resembled living crocodilians, and probably had a similar ecology as semi-aquatic ambush predators.
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". Despite Pseudosuchia meaning "false crocodiles", the name is a misnomer as true crocodilians are now defined as a subset of the group.
Metriorhynchidae is an extinct family of specialized, aquatic metriorhynchoid crocodyliforms from the Middle Jurassic to the Early Cretaceous period of Europe, North America and South America. The name Metriorhynchidae was coined by the Austrian zoologist Leopold Fitzinger in 1843. The group contains two subfamilies, the Metriorhynchinae and the Geosaurinae. They represent the most marine adapted of all archosaurs.
Thalattosuchia is a clade of mostly marine crocodylomorphs from the Early Jurassic to the Early Cretaceous that had a cosmopolitan distribution. They are colloquially referred to as marine crocodiles or sea crocodiles, though they are not members of Crocodilia and records from Thailand and China suggest that some members lived in freshwater. The clade contains two major subgroupings, the Teleosauroidea and Metriorhynchoidea. Teleosauroids are not greatly specialised for oceanic life, with back osteoderms similar to other crocodyliformes. Within Metriorhynchoidea, the Metriorhynchidae displayed extreme adaptions for life in the open ocean, including the transformation of limbs into flippers, the development of a tail fluke, and smooth, scaleless skin, and probably gave live birth, seemingly uniquely among archosaurs.
Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day.
Sphenosuchia is a suborder of basal crocodylomorphs that first appeared in the Triassic and occurred into the Middle Jurassic. Most were small, gracile animals with an erect limb posture. They are now thought to be ancestral to crocodyliforms, a group which includes all living crocodilians.
Terrestrisuchus is an extinct genus of very small early crocodylomorph that was about 76 centimetres (30 in) long. Fossils have been found in Wales and Southern England and date from near the very end of the Late Triassic during the Rhaetian, and it is known by type and only known species T. gracilis. Terrestrisuchus was a long-legged, active predator that lived entirely on land, unlike modern crocodilians. It inhabited a chain of tropical, low-lying islands that made up southern Britain, along with similarly small-sized dinosaurs and abundant rhynchocephalians. Numerous fossils of Terrestrisuchus are known from fissures in limestone karst which made up the islands it lived on, which formed caverns and sinkholes that preserved the remains of Terrestrisuchus and other island-living reptiles.
Pelagosaurus is an extinct genus of thalattosuchian crocodyliform that lived during the Toarcian stage of the Lower Jurassic, around 183 Ma to 176 Ma, in shallow epicontinental seas that covered much of what is now Western Europe. The systematic taxonomy of Pelagosaurus has been fiercely disputed over the years, and was assigned to Thalattosuchia after its systematics within Teleosauridae were disputed. Pelagosaurus measured 2–3 m (6.6–9.8 ft) long.
Junggarsuchus is an extinct genus of sphenosuchian crocodylomorph from the Middle or Late Jurassic period of China. The type and only species is J. sloani. The generic name of Junggarsuchus comes from the Junggar Basin, where the fossil was found, and the Greek word "souchos" meaning crocodile. The specific name, "sloani" is in honor of C. Sloan, who is credited with finding the holotype.
Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group ; if Sebecosuchia is included within Notosuchia its existence is pushed into the Middle Miocene, about 11 million years ago. Fossils have been found from South America, Africa, Asia, and Europe. Notosuchia was a clade of terrestrial crocodilians that evolved a range of feeding behaviours, including herbivory (Chimaerasuchus), omnivory (Simosuchus), and terrestrial hypercarnivory (Baurusuchus). It included many members with highly derived traits unusual for crocodylomorphs, including mammal-like teeth, flexible bands of shield-like body armor similar to those of armadillos (Armadillosuchus), and possibly fleshy cheeks and pig-like snouts (Notosuchus). The suborder was first named in 1971 by Zulma Gasparini and has since undergone many phylogenetic revisions.
Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to Crocodylus niloticus than to Notosuchus terrestris. Members of Neosuchia generally share a crocodilian-like bodyform adapted to freshwater aquatic life, as opposed to the terrestrial habits of more basal crocodylomorph groups. The earliest neosuchian is suggested to be the Early Jurassic Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages in North America. It is often identified as a member of Goniopholididae, though this is disputed, and the taxon may lie outside Neosuchia, which places the earliest records of the group in the Middle Jurassic.
Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudosuchians to survive the K-Pg extinction event.
Pholidosaurus is an extinct genus of neosuchian crocodylomorph. It is the type genus of the family Pholidosauridae. Fossils have been found in northwestern Germany. The genus is known to have existed during the Berriasian-Albian stages of the Early Cretaceous. Fossil material found from the Annero and Jydegård Formations in Skåne, Sweden and on the island of Bornholm, Denmark, have been referred to as a mesoeucrocodylian, and possibly represent the genus Pholidosaurus.
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.
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.
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.
Almadasuchus is an extinct genus of crocodylomorph known from the early Late Jurassic Puesto Almada Member of the e Cañadón Calcáreo Formation of Patagonia, Argentina. It contains a single species, Almadasuchus figarii. It is known from the holotype MPEF-PV 3838, a well-preserved posterior region of the skull as well as other skull and postcranial remains. Almadasuchus was recovered from Puesto Almada, 30 m above the fish beds, dated as Oxfordian in age.