Crurotarsi is a

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

of archosauriform reptiles that includes

crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...

ns and stem-crocodilians and possibly bird-line

archosaurs Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...

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 Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

. An equivalent term for the crocodilian side of the archosaur family tree is

Pseudosuchia Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...

. This traditional definition of Crurotarsi assumed that phytosaurs were

crown-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. ...

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. The name Crurotarsi is derived from the Latin word '' crus'' (lower leg) and the Greek word '' tarsos'' (ankle). It refers to the specialized articulation (a

crurotarsal A crurotarsal joint is one that's situated between the bones of '' crus'', i.e. shin (tibia and fibula) and the proximal tarsal bones, i.e. astragalus and calcaneum. The ankle joint of therian mammals ( marsupials and placentals) is a crurotarsa ...

joint) between the lower leg (specifically the

fibula The fibula (: fibulae or fibulas) 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. ...

) and the ankle (specifically the

calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel; : calcanei or calcanea) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is t ...

) which is present in the skeletons of reptiles such as

suchia Suchia (from Ancient Greek σούχος (''soúkhos''), meaning "crocodile") is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing ''A ...

ns and phytosaurs. In their ankle joint, a hemicylindrical

condyle A condyle (;Entry "condyle"
in

on the calcaneum articulates into a concave area on the fibula.

Taxonomic history

The name Crurotarsi was erected as a node-based clade by

Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago who has discovered several new dinosaur species on several continents, including at sites in Inner Mongolia, Argentina, Morocco and Niger. ...

and A. B. Arcucci in 1990 to supplant the old term

, but with a different definition. Crurotarsi includes, by most published definitions, all descendants of the common ancestor of modern

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 more loosely to include ...

s, ornithosuchids,

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, herbivor ...

s, and phytosaurs; Nesbitt (2011) provided a shorter definition, defining Crurotarsi as "the least inclusive clade containing '' Rutiodon carolinensis'' Emmons, 1856, and ''

Crocodylus niloticus 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 in sub-Saharan Africa, occurring mostly in the eastern, southern, and ce ...

'' Laurenti, 1768". According to two studies published in 2011 by Nesbitt and coworkers, using either of these definitions leads to the inclusion of all other true

in Crurotarsi, due to the possibly basal phylogenetic position of the phytosaurs. This means that grouping the phytosaurs and crocodilians into a clade while excluding the

avemetatarsalia Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid Reptile, reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosau ...

ns (

pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...

s, and

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s) would result in a paraphyletic grouping. A more definitive group is

, which is defined as all archosaurs closer to crocodiles than to birds (matching the traditional content of Crurotarsi).

Phylogeny

Paul Sereno and A. B. Arcucci named Crurotarsi in 1990, defining it as " Parasuchia hytosaurs Ornithosuchidae, '' Prestosuchus'', Suchia, and all descendants of their common ancestor". The groups in this definition were considered crocodile-line archosaurs, as opposed to the bird-line archosaurs. Ornithosuchids were once considered bird-line archosaurs (as implied by their name, which means "bird crocodiles" in

Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...

), but were later recognized as crocodile-line archosaurs. This reclassification may have inspired Sereno's Crurotarsi, a node-based clade defined by the inclusion of ornithosuchids and other early archosaurs. Two names were proposed for crocodile-line archosaurs before Crurotarsi was erected. The first, Pseudosuchia, was established as a stem-based clade in 1985. It includes crocodiles and all archosaurs more closely related to crocodiles than to birds. The second, Crocodylotarsi, was named in 1988, possibly as a replacement for Pseudosuchia. The name Pseudosuchia, meaning "false crocodiles", has been used for over a century, and traditionally included aetosaurs. As a clade, Pseudosuchia includes the group

Eusuchia Eusuchia is a clade of neosuchian Crocodylomorpha, crocodylomorphs that first appeared in the Early Cretaceous, which includes modern Crocodilia, crocodilians. Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survi ...

, or "true crocodiles". Crocodylotarsi may have been named to remove confusion, but as a stem-based clade it is synonymous with Pseudosuchia. Because Pseudosuchia was named first, it has precedence. Crurotarsi traditionally contains the same archosaurs as Pseudosuchia, but as a node-based clade it is not synonymous. Below is a cladogram after Nesbitt & Norell (2006) and Nesbitt (2007) with Crurotarsi in its traditional sense encompassing just crocodile-line archosaurs: Cladogram after Brusatte, Benton, Desojo and Langer (2010): In 2011, Sterling J. Nesbitt found phytosaurs to be the sister taxon of Archosauria, and therefore not crocodile-line archosaurs. Because phytosaurs are included in the definition of Crurotarsi, this change in their

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

placement expanded the scope of Crurotarsi, which therefore now includes phytosaurs, crocodiles, pterosaurs and dinosaurs. However, Pseudosuchia still contains only crocodile-line archosaurs. Below is a

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

modified from Nesbitt (2011) showing the new changes:

References

* * *

External links

Palaeos
{{Taxonbar, from=Q131341 Archosauriformes Tetrapod unranked clades Triassic reptiles Extant Early Triassic first appearances Taxa named by Paul Sereno