Phytosaur in the context of "Archosaurian"

Archosauria (lit. 'ruling reptiles') or archosaurs (/ˈɑːrkəˌsɔːr/) is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only known extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistic sense of the term includes all living and extinct relatives of birds and crocodilians such as non-avian dinosaurs, pterosaurs, phytosaurs, aetosaurs and rauisuchians as well as many Mesozoic marine reptiles. 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 (such as non-avian dinosaurs and pterosaurs). Older definitions of the group Archosauria rely on shared morphological characteristics, such as an antorbital fenestra in the skull, serrated teeth, and an upright stance. Some extinct reptiles, such as proterosuchids and euparkeriids, also possessed these features yet originated prior to the split between the crocodilian and bird lineages. The older morphological definition of Archosauria nowadays roughly corresponds to Archosauriformes, a group named to encompass crown-group archosaurs and their close relatives.

Pseudosuchia (from Ancient Greek ψεύδος (pseúdos), meaning "false", and Σοῦχος (Soûkhos), meaning "Sobek") 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", in contrast to the "bird-line archosaurs" or Avemetatarsalia. Despite Pseudosuchia meaning "false crocodiles", the name is a misnomer as true crocodilians are now defined as a subset of the group.

The clade Pseudosuchia is potentially equivalent to another term, Crurotarsi, even though the latter has a different, node-based definition: "all taxa the least inclusive clade containing Rutiodon carolinensis (Emmons, 1856), and Crocodylus niloticus (Laurenti, 1768)." Many paleontologists of the late 20th century took this proposal for granted, using Crurotarsi as the term for crocodilian ancestors. In 2011, a major revision of Triassic archosaur relations proposed that Rutiodon's group, Phytosauria, was not as closely related to other traditional "crurotarsans", at least compared to avemetatarsalians such as pterosaurs and dinosaurs. Under that interpretation, Crurotarsi would be a much broader clade than Pseudosuchia. Other recent studies continue to support a more traditional phylogeny with phytosaurs as an early branch of Pseudosuchia. If the traditional interpretation is maintained, Pseudosuchia and Crurotarsi are roughly equivalent categories.

"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 (often 4 to 6 metres (13 to 20 ft)), 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 (crocodile-like carnivores), aetosaurs (armored herbivores), and crocodylomorphs (lightly-built crocodilian ancestors).

However, more recent studies on archosaur evolution have upended this idea based on phylogenetic analyses and cladistics, a modern approach to taxonomy based on clades (nested monophyletic groups of common ancestry). Since the early 2010s, archosaur classification schemes have stabilized on a system where Rauisuchia is rendered an evolutionary grade, or even a wastebin taxon. Crocodylomorphs most likely originated from a rauisuchian ancestor based on a myriad of shared traits, and some "rauisuchians" (such as Postosuchus and Rauisuchus) appear to be more closely related to crocodylomorphs than to other "rauisuchians" (such as Prestosuchus and Saurosuchus).

Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amphibians), various groups of dinosaurs (most notably ankylosaurs and stegosaurians), phytosaurs, aetosaurs, placodonts, and hupehsuchians (marine reptiles with possible ichthyosaur affinities).

Osteoderms are uncommon in mammals, although they have occurred in many xenarthrans (armadillos and the extinct glyptodonts and mylodontid ground sloths). The heavy, bony osteoderms have evolved independently in many different lineages. The armadillo osteoderm is believed to develop in subcutaneous dermal tissues. These varied structures should be thought of as anatomical analogues, not homologues, and do not necessarily indicate monophyly. The structures are however derived from scutes, common to all classes of amniotes and are an example of what has been termed deep homology. In many cases, osteoderms may function as defensive armor. Osteoderms are composed of bone tissue, and are derived from a scleroblast neural crest cell population during embryonic development of the organism. The scleroblastic neural crest cell population shares some homologous characteristics associated with the dermis. Neural crest cells, through epithelial-to-mesenchymal transition, are thought to contribute to osteoderm development.

Rutiodon (meaning "wrinkle tooth") is an extinct genus of mystriosuchine phytosaurs from the Late Triassic of the eastern United States. The type species of Rutiodon, Rutiodon carolinensis, encompasses a large number of skulls and assorted postcranial fossils discovered in the Cumnock Formation of North Carolina. Fossils referable to the species are also known from Pennsylvania, New Jersey, and Virginia. Rutiodon carolinensis is the most well-described species of phytosaur in eastern North America, though its validity as a natural taxon has been questioned. Some paleontologists also recognize a larger and more robust species, Rutiodon manhattanensis, which is known from teeth and postcranial fossils from New Jersey and Pennsylvania.

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.

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 joint) between the lower leg (specifically the fibula) and the ankle (specifically the calcaneum) which is present in the skeletons of reptiles such as suchians and phytosaurs. In their ankle joint, a hemicylindrical condyle on the calcaneum articulates into a concave area on the fibula.

Parasuchidae is a clade of phytosaurs more derived than Diandongosuchus, a basal phytosaur. This family was phylogenetically defined by Christian Kammerer and colleagues in 2015 as the last common ancestor and all descendants of Wannia scurriensis, Parasuchus hislopi, and Mystriosuchus planirostris. It encompasses nearly all phytosaurs, including early Parasuchus-grade forms as well as a more restricted clade of more specialized phytosaurs. This more restricted clade is traditionally known as the family Phytosauridae and more recently as the subfamily Mystriosuchinae, defined by Kammerer et al. 2015 as the last common ancestor and all descendants of Mystriosuchus planirostris and Angistorhinus grandis.

Parasuchids have been recovered from Late Triassic deposits in Europe, North America, India, Morocco, Thailand, Brazil, Greenland and Madagascar. In their osteology of Parasuchus, Kammerer et al. (2016) suggested using Parasuchidae to include taxa traditionally included in Phytosauridae as well as Parasuchus-grade taxa. Stocker et al. (2017) use the phytosaur classification advocated by Kammerer et al. (2016) by recovering Diandongosuchus as the basalmost phytosaur outside Parasuchidae, noting that Diandongosuchus has a shorter snout than parasuchids.