Crocodylomorpha is a group 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 that includes the

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 and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During

Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...

and early Cenozoic times, crocodylomorphs were far more diverse than they are now. Triassic forms were small, lightly built, active terrestrial animals. The earliest and most primitive crocodylomorphs are represented by " sphenosuchians", a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

assemblage containing small-bodied forms with elongated limbs that walked upright, which represents the ancestral morphology of Crocodylomorpha. These forms persisted until the end of the Jurassic. During the Jurassic, Crocodylomorphs morphologically diversified into numerous niches, including into the aquatic and marine realms.

Evolutionary history

When their extinct species and

stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

are examined, the crocodylian lineage (clade

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 ...

, formerly

Crurotarsi 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 though ...

) proves to have been a very diverse and adaptive group of reptiles. Not only are they an ancient group of animals – at least as old as the

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 ...

s – they also evolved into a great variety of forms and sizes. Some terrestrial

notosuchia 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 (see below); if Sebecosu ...

ns were noticeably small animals, less than in length, while aquatic or semiaquatic crocodylomorphs, especially tethysuchians,

thalattosuchia Thalattosuchia is a clade of 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 Cro ...

ns, and some

ns reached around in length. The earliest forms, the

sphenosuchia 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, ...

ns, evolved during the Late

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 ...

, and were highly gracile terrestrial forms built like greyhounds. During the

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...

and the Cretaceous, marine forms in the family Metriorhynchidae, such as ''

Metriorhynchus ''Metriorhynchus'' is an extinct genus of marine crocodyliform that lived in the oceans during the Late Jurassic. The type species, ''M. brevirostris'' was named in 1829 as a species of ''Steneosaurus'' before being named as a separate genus by ...

'', evolved forelimbs that were paddle-like and had a tail similar to modern fish. '' Dakosaurus andiniensis'', a species closely related to ''Metriorhynchus'', had a skull that was adapted to eat large marine reptiles. Several terrestrial species during the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

were herbivorous, such as '' Simosuchus clarki'' and ''

Chimaerasuchus paradoxus ''Chimaerasuchus'' (" chimera crocodile") is an extinct genus of Chinese crocodile like form from the Early Cretaceous Wulong Formation. The four teeth in the very tip of its short snout gave it a "bucktoothed" appearance. Due its multicusped ...

''. A number of lineages during the Cenozoic became wholly terrestrial predators.

Taxonomy and phylogeny

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 ''Mekosuchus'' is a genus of extinct Australasian crocodiles within the subfamily Mekosuchinae. They are believed to have been made extinct by the arrival of humans on the South Pacific islands where they lived. The species of this genus were s ...

''. The various other groups that had previously been known as Crocodilia were moved to Crocodylomorpha and the slightly more restrictive

Crocodyliformes 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 pse ...

. Crocodylomorpha has been given the rank of

superorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

in some 20th and 21st century studies. The old Crocodilia was subdivided into the suborders: *

Eusuchia Eusuchia is a clade of crocodylomorphs that first appeared in the Early Cretaceous with '' Hylaeochampsa''. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-T extinction. Since the other two clades ...

: true crocodiles (which includes crown-group

Crocodylia 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 ...

) * Mesosuchia: 'middle' crocodiles *

Thalattosuchia Thalattosuchia is a clade of 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 Cro ...

: sea crocodiles *

Protosuchia Protosuchia is a group of extinct Mesozoic crocodyliforms. They were small in size (~1 meter in length) and terrestrial. In phylogenetic terms, Protosuchia is considered an informal group because it is a grade of basal crocodyliforms, not a tru ...

: first crocodiles Mesosuchia is a

group as it does not include eusuchians (which nest within Mesosuchia).

Mesoeucrocodylia 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. Diagnosis It was long known that M ...

was the name given to the clade that contains mesosuchians and eusuchians (Whetstone and Whybrow, 1983).

Phylogeny

Below is a

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

modified from Nesbitt (2011) and Bronzati (2012). 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).

Biology

The Crocodylomorpha comprise a variety of forms, shapes, and sizes, which occupied a range of habitats. As with most

amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...

s, Crocodylomorphs were and are

oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and ...

, laying eggs in a nest or mound, known from strata as old as the Late Jurassic. Adult size varies widely, from about 55 cm long in ''

Knoetschkesuchus ''Knoetschkesuchus'' is a genus of small atoposaurid crocodylomorph from the Late Jurassic of Germany and Portugal. Two species are known: the German species ''K. langenbergensis'', described by Schwarz and colleagues in 2017 based on two part ...

'' to much larger dimensions, as in ''

Sarcosuchus ''Sarcosuchus'' (; ) is an extinct genus of crocodyliform and distant relative of living crocodilians that lived during the Early Cretaceous, from the late Hauterivian to the early Albian, 133 to 112 million years ago of what is now Africa and ...

''. Most crocodylomorphs were

carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...

s, but many lineages evolved to be obligate

piscivore A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evoluti ...

s, such as the extant gharials.

References

Sources

* Benton, M. J. (2004), ''

Vertebrate Palaeontology Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct animals with vertebrae or a notochord. It also tries to connect, by u ...

'', 3rd ed. Blackwell Science Ltd * Hay, O. P. 1930 (1929–1930). Second Bibliography and Catalogue of the Fossil Vertebrata of North America. ''Carnegie Institution Publications'', Washington, 1, 990 pp.

External links

Crocodylomorpha
- webpages by Ross Elgin on the

University of Bristol , mottoeng = earningpromotes one's innate power (from Horace, ''Ode 4.4'') , established = 1595 – Merchant Venturers School1876 – University College, Bristol1909 – received royal charter , type ...

server *
Major subgroups
classification (used here)

from

Palaeos Palaeos.com is a web site on biology, paleontology, phylogeny and geology and which covers the history of Earth. The site is well respected and has been used as a reference by professional paleontologists such as Michael J. Benton, the professor of ...

*
Technical definition
{{Taxonbar, from=Q131863 01 Chordate superorders Extant Late Triassic first appearances Norian first appearances