Eusuchia

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Eusuchia
Temporal range: Early Cretaceous - Recent, 130–0  Ma
Comparison - Crocodilia.jpg
Three species of living eusuchian: gharial (left), American alligator (center), and American crocodile (right).
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Pseudosuchia
Clade: Crocodylomorpha
Clade: Crocodyliformes
Clade: Neosuchia
Clade: Eusuchia
Huxley, 1875
Subgroups

Eusuchia is a clade of neosuchian crocodylomorphs that first appeared in the Early Cretaceous, which includes modern crocodilians. [3] Along with Dyrosauridae and Sebecosuchia, they were the only crocodyliformes who survived the K-Pg extinction.

Contents

Definition

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". [4] 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.

Description

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. [5]

Phylogeny

A 2018 tip dating study by Lee & Yates using both molecular, morphological and stratigraphic data recovered the cladogram below: [2]

A 2021 Rio & Mannion phylogenetic study using a heavily modified morphological data set recovered the cladogram below: [6]

Related Research Articles

<span class="mw-page-title-main">Gavialinae</span> Subfamily of gharial crocodylians

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.

<span class="mw-page-title-main">Gavialidae</span> Family of gharial crocodylians

Gavialidae is a family of large semiaquatic crocodilians with elongated, narrow snouts. Gavialidae consists of two living species, the gharial and the false gharial, both occurring in Asia. Many extinct members are known from a broader range, including the recently extinct Hanyusuchus. Gavialids are generally regarded as lacking the jaw strength to capture the large mammalian prey favoured by crocodiles and alligators of similar size so their thin snout is best used to catch fish, however the false gharial has been found to have a generalist diet with mature adults preying upon larger vertebrates, such as ungulates.

<i>Allodaposuchus</i> Extinct genus of reptiles

Allodaposuchus is an extinct genus of crocodyliforms that lived in what is now southern Europe during the Campanian and Maastrichtian stages 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.

<span class="mw-page-title-main">Crocodylomorpha</span> Clade of reptiles

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. Extinct crocodylomorphs were considerably more ecologically diverse tham modern crocodillians. The earliest and most primitive crocodylomorphs are represented by "sphenosuchians", a paraphyletic assemblage containing small-bodied, slender 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, with the subgroups Neosuchia and the extinct Thalattosuchia adapting to aquatic life, while some terrestrial groups adopted herbivorous and omnivorous lifestyles. Terrestrial crocodylomorphs would continue to co-exist alongside aquatic forms until becoming extinct during the Miocene.

<i>Boverisuchus</i> Extinct genus of reptiles

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).

<i>Borealosuchus</i> Extinct genus of reptiles

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.

Argochampsa is an extinct genus of eusuchian crocodylomorph, usually regarded as a gavialoid crocodilian, related to modern gharials. It lived in the Paleocene of Morocco. Described by Hua and Jouve in 2004, the type species is A. krebsi, with the species named for Bernard Krebs. Argochampsa had a long narrow snout, and appears to have been marine in habits.

<span class="mw-page-title-main">Alligatoroidea</span> Superfamily of reptiles

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.

<i>Eothoracosaurus</i> Extinct genus of reptiles

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.

<i>Thoracosaurus</i> Extinct genus of reptiles

Thoracosaurus is an extinct genus of long-snouted eusuchian which existed during the Late Cretaceous and Early Paleocene in North America and Europe.

<i>Planocrania</i> Extinct genus of reptiles

Planocrania is an extinct genus of eusuchian crocodyliforms from what is now China. Two species are currently known to belong to the genus.

<span class="mw-page-title-main">Crocodyloidea</span> Superfamily of crocodiles

Crocodyloidea is one of three superfamilies of crocodilians, the other two being Alligatoroidea and Gavialoidea, and it includes the crocodiles. Crocodyloidea may also include the extinct Mekosuchinae, native to Australasia from the Eocene to the Holocene, although this is disputed.

<span class="mw-page-title-main">Gavialoidea</span> Superfamily of large reptiles

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.

<span class="mw-page-title-main">Brevirostres</span> Taxon of reptiles

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:

<span class="mw-page-title-main">Planocraniidae</span> Extinct family of reptiles

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".

<span class="mw-page-title-main">Allodaposuchidae</span> Extinct clade of reptiles


Allodaposuchidae is an extinct clade of eusuchians that lived in Europe during the Late Cretaceous (Santonian-Maastrichtian).

<i>Lohuecosuchus</i> Extinct genus of reptiles

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:

<span class="mw-page-title-main">Longirostres</span> Clade of crocodilians

Longirostres is a clade of crocodilians that includes the crocodiles and the gavialids, to the exclusion of the alligatoroids. Defined in 2003 by Harshman et al., Longirostres is a crown group defined phylogenetically as including the last common ancestor of Crocodylus niloticus and Gavialis gangeticus and all of its descendants.

References

  1. Casey M. Holliday & Nicholas M. Gardner (2012). "A New Eusuchian Crocodyliform with Novel Cranial Integument and Its Significance for the Origin and Evolution of Crocodylia". PLOS ONE. 7 (1): e30471. Bibcode:2012PLoSO...730471H. doi: 10.1371/journal.pone.0030471 . PMC   3269432 . PMID   22303441. Open Access logo PLoS transparent.svg
  2. 1 2 Michael S. Y. Lee; Adam M. Yates (27 June 2018). "Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil". Proceedings of the Royal Society B . 285 (1881). doi: 10.1098/rspb.2018.1071 . PMC   6030529 . PMID   30051855.
  3. Benton, Michael J.; Sibbick, John (2000). Vertebrate Palaeontology. Blackwell Publishing. p. 233. ISBN   0-632-05614-2.
  4. Brochu, Christopher A. (2003). "PHYLOGENETIC APPROACHES TOWARD CROCODYLIAN HISTORY". Annual Review of Earth and Planetary Sciences (31): 357–97. doi:10.1146/annurev.earth.31.100901.141308.
  5. Puértolas-Pascual, E; Mateus, O (2020-06-11). "A three-dimensional skeleton of Goniopholididae from the Late Jurassic of Portugal: implications for the Crocodylomorpha bracing system". Zoological Journal of the Linnean Society. 189 (2): 521–548. doi:10.1093/zoolinnean/zlz102. ISSN   0024-4082.
  6. Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ . 9: e12094. doi: 10.7717/peerj.12094 . PMC   8428266 . PMID   34567843.