Eusuchia Temporal range: Early Cretaceous - Recent, | |||
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Three species of living eusuchian: gharial (left), American alligator (center), and American crocodile (right). | |||
Scientific classification | |||
Domain: | Eukaryota | ||
Kingdom: | Animalia | ||
Phylum: | Chordata | ||
Class: | Reptilia | ||
Clade: | Archosauria | ||
Clade: | Pseudosuchia | ||
Clade: | Crocodylomorpha | ||
Clade: | Crocodyliformes | ||
Clade: | Neosuchia | ||
Clade: | Eusuchia Huxley, 1875 | ||
Subgroups | |||
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Eusuchia is a clade of neosuchian crocodylomorphs that first appeared in the Early Cretaceous, which includes modern crocodilians. [8] Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-Pg extinction.
Eusuchia was originally defined by Thomas Henry Huxley in 1875 as an apomorphy-based group, meaning that it was defined by shared characteristics rather than relations. These characteristics include pterygoid-bounded choanae and vertebrae which are procoelous (concave from the front and convex from the back). The possibility that these traits may have been convergently evolved in different groups of neosuchians rather than one lineage spurred some modern paleontologists to revise the group's definition to make it defined solely by relations. In 1999, Christopher Brochu redefined Eusuchia as "the last common ancestor of Hylaeochampsa and Crocodylia and all of its descendants". [9] In this definition, "Crocodylia" specifically refers to descendants of the common ancestor of the three modern lineages of eusuchians: Gavialoidea (gharials), Alligatoroidea (alligators and caimans), and Crocodyloidea (crocodiles). Whether certain families or genera qualify as basal eusuchians or non-eusuchian neosuchians is a matter of debate over which definition is used for the group. At least one genus, Isisfordia , would qualify as a eusuchian under Huxley's definition of the group but would not necessarily qualify under Brochu's definition.
Unlike primitive crocodylomorphs, crocodyliforms have secondary bony palates. This feature enables living crocodylians to safely breathe in through their nostrils while the rest of the head (including the mouth) remains submerged. This structure reaches its greatest elaboration among eusuchians, in which the internal nares are completely surrounded by the pterygoid bones.
Unlike more basal neosuchians, eusuchians have four rather than two rows of dorsal armour plates, which are square in shape, with 2 to 4 rows of accessory plates. [10]
A 2018 tip dating study by Lee & Yates using both molecular, morphological and stratigraphic data recovered the cladogram below: [3]
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A 2021 Rio & Mannion phylogenetic study using a heavily modified morphological data set recovered the cladogram below: [11]
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Gavialinae is a subfamily of large semiaquatic crocodilian reptiles, resembling crocodiles, but with much thinner snouts. Gavialinae is one of the two major subfamilies within the family Gavialidae - the other being the subfamily Tomistominae, which contains the false gharial and extinct relatives.
Allodaposuchus is an extinct genus of crocodyliforms that lived in what is now southern Europe during the Campanian and Maastrichtian stages, and possibly the Santonian stage, of the Late Cretaceous. Although generally classified as a non-crocodylian eusuchian crocodylomorph, it is sometimes placed as one of the earliest true crocodylians. Allodaposuchus is one of the most common Late Cretaceous crocodylomorphs from Europe, with fossils known from Romania, Spain, and France.
Boverisuchus is an extinct genus of planocraniid crocodyliforms known from the middle Eocene of Germany and western North America. It was a relatively small crocodyliform with an estimated total length of approximately 2.2–3.6 metres (7.2–11.8 ft).
Borealosuchus is an extinct genus of crocodyliforms that lived from the Late Cretaceous to the Eocene in North America. It was named by Christopher Brochu in 1997 for several species that had been assigned to Leidyosuchus. The species assigned to it are: B. sternbergii, the type species, from the Maastrichtian of Colorado, Montana, North Dakota, South Dakota, and Wyoming; B. acutidentatus, from the Paleocene of Saskatchewan; B. formidabilis, from the Paleocene of North Dakota; B. griffithi, from the Paleocene of Alberta; and B. wilsoni, from the Eocene of Wyoming. B. formidabilis is particularly well-known, represented by the remains of many individuals from the Wannagan Creek site in North Dakota. An indeterminate species is known from the Late Cretaceous Demopolis Chalk in Alabama.
Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more closely related to the alligators than the two other groups.
Iharkutosuchus is an extinct genus of basal eusuchian crocodyliform. Its fossils have been found in the Santonian-age Upper Cretaceous Csehbánya Formation in the Bakony Mountains of western Hungary.
Isisfordia is an extinct genus of crocodyliform closely related to crocodilians that lived in Australia during the Middle Cretaceous (Albian–Cenomanian).
Eothoracosaurus is an extinct monospecific genus of eusuchian crocodylomorphs found in Eastern United States which existed during the Late Cretaceous period. Eothoracosaurus is considered to belong to an informally named clade called the "thoracosaurs", named after the closely related Thoracosaurus. Thoracosaurs in general were traditionally thought to be related to the modern false gharial, largely because the nasal bones contact the premaxillae, but phylogenetic work starting in the 1990s instead supported affinities within gavialoid exclusive of such forms. Even more recent phylogenetic studies suggest that thoracosaurs might instead be non-crocodilian eusuchians.
Hylaeochampsa is an extinct genus of eusuchian crocodylomorphs. It is known only from a partial skull recovered from Barremian-age rocks of the Lower Cretaceous Vectis Formation of the Isle of Wight. This skull, BMNH R 177, is short and wide, with a eusuchian-like palate and inferred enlarged posterior teeth that would have been suitable for crushing. Hylaochampsa was described by Richard Owen in 1874, with H. vectiana as the type species. It may be the same genus as the slightly older Heterosuchus, inferred to have been of similar evolutionary grade, but there is no overlapping material as Heterosuchus is known only from vertebrae. If the two could be shown to be synonyms, Hylaeochampsa would have priority because it is the older name. Hylaeochampsa is the type genus of the family Hylaeochampsidae, which also includes Iharkutosuchus from the Late Cretaceous of Hungary. James Clark and Mark Norell positioned it as the sister group to Crocodylia. Hylaeochampsa is currently the oldest known unambiguous eusuchian.
Hylaeochampsidae is an extinct family of basal eusuchian crocodylomorphs thought to be closely related to the order Crocodylia.
Thoracosaurus is an extinct genus of long-snouted eusuchian which existed during the Late Cretaceous and Early Paleocene in North America and Europe.
Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.
Stangerochampsa is an extinct genus of alligatorid, possibly an alligatorine or a stem-caiman, from the Late Cretaceous of Alberta. It is based on RTMP.86.61.1, a skull, partial lower jaws, and partial postcranial skeleton discovered in the late Campanian–early Maastrichtian-age Horseshoe Canyon Formation. Stangerochampsa was described in 1996 by Wu and colleagues. The type species is S. mccabei. The generic name honors the Stanger family, the owners of the ranch where the specimen was found, and the species name honors James Ross McCabe, who discovered, collected, and prepared it. Stangerochampsa is described as "small to medium–sized"; the type skull is 20.0 centimetres (7.9 in) long from the tip of the snout to the occipital condyle, and is 13.0 centimetres (5.1 in) wide at its greatest, while the thigh bone is 14.2 centimetres (5.6 in) long. It had heterodont dentition, with large crushing teeth at the rear of the jaws.
Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial Gavialis gangeticus and the false gharial Tomistoma schlegelii are alive today, with Hanyusuchus having become extinct in the last few centuries.
Brevirostres is a paraphyletic group of crocodilians that included alligatoroids and crocodyloids. Brevirostres are crocodilians with small snouts, and are distinguished from the long-snouted gharials. It is defined phylogenetically as the last common ancestor of Alligator mississippiensis and Crocodylus niloticus and all of its descendants. This classification was based on morphological studies primarily focused on analyzing skeletal traits of living and extinct fossil species, and placed the gharials outside the group due to their unique skull structure, and can be shown in the simplified cladogram below:
Planocraniidae is an extinct family of eusuchian crocodyliforms known from the Paleogene of Asia, Europe and North America. The family was coined by Li in 1976, and contains three genera, Boverisuchus, Duerosuchus and Planocrania. Planocraniids were highly specialized crocodyliforms that were adapted to living on land. They had extensive body armor, long legs, and blunt claws resembling hooves, and are sometimes informally called "hoofed crocodiles".
Allodaposuchidae is an extinct clade of eusuchians that lived in Europe during the Late Cretaceous (Santonian-Maastrichtian).
Lohuecosuchus is an extinct genus of allodaposuchid eusuchian crocodylomorph that lived during the Late Cretaceous in what is now Spain and southern France.
Portugalosuchus is an extinct genus of eusuchian crocodyliform that was possibly a basal crocodylian – if so then it would be the oldest known crocodylian to date. The type species is P. azenhae, described in 2018, and it is known from the Late Cretaceous (Cenomanian)-aged Tentugal Formation in Portugal. A 2021 morphological study recovered Portugalosuchus within Crocodylia as a member of Gavialidae closely related to similar "thoracosaurs", while also noting that it might also possibly be outside of Crocodylia completely. A 2022 tip dating analysis incorporating both morphological and DNA data placed Portugalosuchus outside of Crocodylia, as the sister taxon of the family Allodaposuchidae. A cladogram simplified after that analysis is shown below:
Scolomastax is an extinct genus of paralligatorid neosuchian known from the Late Cretaceous Woodbine Formation in Texas. It contains a single species, S. sahlsteini.