Crocodyliformes is a
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, ...
of
crurotarsan
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 thoug ...
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, the group often traditionally referred to as "crocodilians". They are the first members of
Crocodylomorpha
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.
During Mesozoic and early Cenozoic times, cro ...
to possess many of the features that define later relatives. They are the only
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 ...
ns to survive the
K-Pg extinction event.
In 1988,
Michael J. Benton
Michael James Benton One or more of the preceding sentences incorporates text from the royalsociety.org website where: (born 8 April 1956) is a British palaeontologist, and professor of vertebrate palaeontology in the School of Earth Sciences ...
and
James M. Clark argued that all traditional names for well-known groups of animals should be restricted to their
crown clade
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. ...
s, that is, used only for natural groups comprising all living members of any given lineage. This posed a problem for the crocodilians, because the name
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 ...
, while used in various ways by various scientists, had always included not only living crocodilians but many of their extinct ancestors known only from the fossil record.
[Benton, M.J. and Clark, J.M. (1988). "Archosaur phylogeny and the relationships of the Crocodylia." Pp. 295–338 in Benton, M.J. (ed.), ''The phylogeny and classification of the Tetrapods, volume 1''. Oxford: Clarendon Press.]
Benton and Clark's solution to this issue was to restrict the name Crocodylia to the group containing modern
alligator
An alligator is a large reptile in the Crocodilia order in the genus ''Alligator'' of the family Alligatoridae. The two extant species are the American alligator (''A. mississippiensis'') and the Chinese alligator (''A. sinensis''). Additiona ...
s,
crocodile
Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s, and
gharial
The gharial (''Gavialis gangeticus''), also known as gavial or fish-eating crocodile, is a crocodilian in the family Gavialidae and among the longest of all living crocodilians. Mature females are long, and males . Adult males have a distinct b ...
s, plus any extinct members of those specific families. The traditional group "Crocodylia" was replaced by the name Crocodyliformes, which included many of the extinct families that the new definition left out. Clark and Benton did not initially provide an exact definition for Crocodyliformes; but, in 2001,
Paul Sereno
Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
and colleagues defined it as the clade including ''
Protosuchus richardsoni'' and the
Nile crocodile
The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed throughout sub-Saharan Africa, occurring mostly in the central, eastern ...
, plus all descendants of their common ancestor.
Chris Brochu agreed with the assessment that Crocodylia as a name has never had stable contents, and that a series of clades larger than the crown group Crocodylia (including Crocodyliformes and the slightly more inclusive clade
Crocodylomorpha
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.
During Mesozoic and early Cenozoic times, cro ...
) was a good solution.
However, in a 2008 paper, Jeremy Martin and Benton reversed the previous opinion (co-authored by Benton) that Crocodylia should be restricted to the crown group, suggesting that Crocodyliformes should be considered a synonym of a more inclusive Crocodylia, and thus replaced.
Brochu and colleagues rejected this proposal, arguing that the crown definition of Crocodylia is the standard meaning both within and beyond the crocodyliform systematics community.
Ecology
While all extant crocodilians are
carnivorous
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 ...
, the ecological roles of Mesozoic crocodyliforms were more diverse, and included
omnivory
An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
and
herbivory
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
. Herbivorous forms were present from the early Jurassic to the late Cretaceous and are thought to have evolved at least three times.
Fossil tracks
Tracks of a crocodyliform are known since Cretaceous, at least. Tracks representing the ichnofamily
Batrachopodidae are described from the
Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
(late
Aptian
The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
)
Calonda Formation (
Angola
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) by
Mateus ''et al.'' (2017), who name a new ichnotaxon ''
Angolaichnus adamanticus''.
Phylogeny
Below is a simplified
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 ...
based on Fiorelli and Calvo (2007).
In 2012, paleontologists Mario Bronzati, Felipe Chinaglia Montefeltro, and Max C. Langer conducted a broad phylogenetic analysis to produce
supertrees of Crocodyliformes, including 184 species. The
most parsimonious trees were highly resolved, meaning the phylogenetic relationships found in the analysis were highly likely. Below is a consensus tree from the study:
References
{{Taxonbar, from=Q3003172