Pholidosauridae

Pholidosauridae is an extinct family of aquatic neosuchian mesoeucrocodylian crocodylomorphs. Fossils have been found in Europe (Denmark, England, France, Germany, Spain and Sweden), Africa (Algeria, Niger, Mali, Morocco and Tunisia), North America (Canada and the United States) and South America (Brazil and Uruguay). The pholidosaurids first appeared in the fossil record during the Bathonian stage of the Middle Jurassic and became extinct during the Late Turonian stage of the Late Cretaceous.[1]

Sarcosuchus is one of the best known pholidosaurs. It is believed to have attained lengths of up to 12 metres (39 ft 4 in) and weighed up to 8 tonnes (7.9 long tons; 8.8 short tons). One genus, Suchosaurus, once thought to be a pholidosaur,[2] has since been shown to be a spinosaurid theropod dinosaur.[3]

Pholidosauridae
Temporal range: Middle Jurassic - Late Cretaceous, 167.7–89.3 Ma
Pholidosaurus
Pholidosaurus meyeri fossil at the Museum für Naturkunde, Berlin
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Suborder: Tethysuchia
Family: Pholidosauridae
Zittel and Eastman, 1902
Subgroups

See text.

Phylogeny

Pholidosauridae is usually considered to be most closely related to the Dyrosauridae. However, the relationship between these families is not fully understood. Pholidosauridae might be monophyletic,[1] paraphyletic[4] or even a polyphyletic[5] in relation to Dyrosauridae. For example, Fortier, Perea & Schultz (2011) found the family to be monophyletic, and include to main lineages: the ElosuchusMeridiosaurus lineage and the Pholidosaurus lineage. The cladogram below shows their phylogenetic analysis, which is based on an expanded version of Pol and Gasparini (2009) analysis.[1]

Neosuchia
Atoposauridae

Alligatorium

Theriosuchus pusillus

Goniopholididae

Bernissartia fagesii

Eusuchia

Thalattosuchia

Pholidosauridae

Pholidosaurus purbeckensis

Sarcosuchus imperator

Terminonaris robusta

Oceanosuchus boecensis

Elosuchus cherifiensis

Meridiosaurus vallisparadisi

Dyrosauridae

Sokotosuchus ianwilsoni

Dyrosaurus phosphaticus

Hyposaurus rogersii

Rhabdognathus

de Andrade et al. (2011) recovered a paraphyletic traditional Pholidosauridae. In their analysis the "Elosuchus lineage" was found to be basal to the "Pholidosaurus lineage"+Dyrosauridae. They used the name Elosuchidae for the Elosuchus lineage and restricted Pholidosauridae to its type genus. The following cladogram simplified after their analysis.[4]

Neosuchia
Atoposauridae

Theriosuchus pusillus

Theriosuchus guimarotae

Rugosuchus

Bernissartia

Eusuchia

Stolokrosuchus

Goniopholididae

Thalattosuchia

Tethysuchia
Elosuchidae

Sarcosuchus hartti

Sarcosuchus imperator

Vectisuchus

Elosuchus

Pholidosauridae

Pholidosaurus schaumburgensis

Dyrosauridae

Congosaurus

Guarinisuchus

Dyrosaurus maghribensis

Dyrosaurus phosphaticus

References

  1. ^ a b c Daniel Fortier, Daniel Perea & Cesar Schultz (2011). "Redescription and phylogenetic relationships of Meridiosaurus vallisparadisi, a pholidosaurid from the Late Jurassic of Uruguay". Zoological Journal of the Linnean Society. 163 (s1): S66–S108. doi:10.1111/j.1096-3642.2011.00709.x.
  2. ^ Carroll, R.L. 1988. Vertebrate Paleontology and Evolution. WH Freeman and Company, New York ISBN 0-7167-1822-7
  3. ^ Buffetaut, E. (2007). "The spinosaurid dinosaur Baryonyx (Saurischia, Theropoda) in the Early Cretaceous of Portugal." Geological Magazine, 144(6): 1021-1025.
  4. ^ a b Marco Brandalise de Andrade; Richard Edmonds; Michael J. Benton; Remmert Schouten (2011). "A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus". Zoological Journal of the Linnean Society. 163 (s1): S66–S108. doi:10.1111/j.1096-3642.2011.00709.x.
  5. ^ Bronzati, M.; Montefeltro, F. C.; Langer, M. C. (2012). "A species-level supertree of Crocodyliformes". Historical Biology. 24 (6): 598–606. doi:10.1080/08912963.2012.662680.

External links

Aegyptosuchidae

Aegyptosuchidae is an extinct family of eusuchian crocodyliforms from the Cretaceous period of Africa. They are characterized by their large size and flat heads. The family includes two genera, Aegyptosuchus and Aegisuchus.

Anglosuchus

Anglosuchus is an extinct genus of pholidosaurid mesoeucrocodylian. Both species of Anglosuchus were originally assigned to the genus Steneosaurus by Richard Owen in 1884, but were later placed in the new genus. It was once thought to be a teleosaurid but later reassigned to the family Pholidosauridae.

Chalawan (genus)

Chalawan (from Thai: ชาละวัน [t͡ɕʰāːlāwān]) is an extinct genus of pholidosaurid mesoeucrocodylian known from the Early Cretaceous Phu Kradung Formation of Nong Bua Lamphu Province, northeastern Thailand. It contains a single species, Chalawan thailandicus.

Coelognathosuchia

Coelognathosuchia is an extinct clade of neosuchian crocodyliforms that includes Goniopholididae and Pholidosauridae, two families of superficially crocodile-like aquatic crocodyliforms from the Mesozoic. Martin et al. (2014) named the clade after finding goniopholidids and pholidosaurids to group together in their phylogenetic analysis of crocodyliform evolutionary relationships. In their analysis, Pholidosauridae was monophyletic and Goniopholididae was paraphyletic, being an assemblage of successively more basal taxa within Coelognathosuchia. Coelognathosuchia itself was positioned near the base of the larger clade Neosuchia as the sister group to a clade containing the Early Cretaceous neosuchian Bernissartia and Eusuchia, the group that includes all modern crocodilians and their closest extinct relatives.Martin et al. named Coelognathosuchia from the Greek κοῖλος (koĩlos, "concave"), γνάθος (gnáthos, "jaw") and σοῦχος (soũchos, "crocodile"), after a small depression on the surface of the skull between the maxilla and jugal bones in both goniopholidids and pholidosaurs. Other diagonostic features of Coelognathosuchia include orbits (eye sockets) that are narrower than the frontal bone that separates them and smaller than the supratemporal fenestrae (two holes at the back of the skull roof) behind them, a notch between the premaxilla and maxilla at the tip of the snout, and the reduction or absence of the antorbital fenestrae (a pair of holes in the snout in front of each orbit).A close relationship between Goniopholididae and Pholidosauridae conflicts with the hypothesis that pholidosaurids are instead more closely related to the family Dyrosauridae. This alternate phylogeny was found in many analyses, including Jouve et al. (2006), Pol and Gasparini (2009), and de Andrade et al. (2011), who named the clade Tethysuchia. In both studies, Goniopholididae was found to be a more distantly related clade within Neosuchia. In their description of Pholidosaurus specimens from southwestern France, Martin and colleagues reiterated their opinion that Dyrosauridae is not as closely related to Pholidosauridae as stated by previous authors by pointing to numerous shared characters between the Cherves-de-Cognac pholidosaurid remains and Goniopholididae.

Coelosuchus

Coelosuchus is an extinct genus of goniopholidid mesoeucrocodylian. Fossils have been found from the Graneros Shale of the Benton Group in Wyoming, and are of Cenomanian age. It was slightly over 1 meter in length.

Elosuchus

Elosuchus is an extinct genus of neosuchian crocodyliform that lived during the Middle Cretaceous of what is now North Africa (Morocco and Algeria).

Meridiosaurus

Meridiosaurus is an extinct genus of mesoeucrocodylian that is a possible member of the family Pholidosauridae. Remains have been found in the Late Jurassic Tacuarembó Formation in Tacuarembó, Uruguay. The genus was described in 1980 on the basis of a partial rostrum that included the premaxillae and most of the maxillae. The assignment to Pholidosauridae is considered doubtful by some authors, but a 2011 redescription and phylogenetic analysis confirmed the pholidosaurid classification of Meridiosaurus.

Mesosuchia

"Mesosuchia" is an obsolete name for a group of terrestrial, semi-aquatic, or fully aquatic crocodylomorph reptiles. The marine crocodile Metriorhynchus had paddle-like forelimbs, Dakosaurus andiniensis had a skull that was adapted to eat large sea reptiles, and Shamosuchus was adapted for eating molluscs and gastropods. Shamosuchus also looked like modern crocodiles and was very closely related to their direct ancestor.

The "Mesosuchia" were formerly placed at Suborder rank as within Crocodylia. The "first" crocodiles were placed within their own suborder, Protosuchia; whilst extant species where placed within Suborder Eusuchia (meaning 'true crocodiles'). Mesosuchia were the crocodylians "in between". As it is a paraphyletic group however, it is not considered valid anymore. It is replaced by its phylogenetic equivalent Mesoeucrocodylia, which contains the taxa herein, the Crocodylia, and some allied forms more recently discovered.

The "Mesosuchia" was composed as:

Family Hsisosuchidae

Family Gobiosuchidae

Infraorder Notosuchia

Family Notosuchidae

Family Sebecidae

Family Baurusuchidae

Infraorder Neosuchia

Family Trematochampsidae

Family Peirosauridae

Genus Lomasuchus

Genus Montealtosuchus

Family Elosuchidae

Family Atoposauridae

Family Dyrosauridae

Family Pholidosauridae

Genus Sarcosuchus

Infraorder Thalattosuchia - Sea "Crocodiles"

Family Teleosauridae

Family Metriorhynchidae

Genus Dakosaurus

Family Goniopholididae

Family Paralligatoridae

Genus Shamosuchus

Nannosuchus

Nannosuchus (meaning "dwarf crocodile") is an extinct genus of goniopholidid mesoeucrocodylian from the Berriasian of England.

Neosuchia

Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to Crocodylus niloticus (the Nile Crocodile) than to Notosuchus terrestris. Neosuchia is very diverse and may be polyphyletic, as the clade has undergone many revisions since it was first named in 1988. Neosuchians first appear in the Early Jurassic with the earliest known goniopholid Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages.

Oceanosuchus

Oceanosuchus is a genus of pholidosaurid mesoeucrocodylian, a type of marine crocodylomorph. It is known from a skull and partial skeleton found in early Cenomanian-age rocks from Normandy, France. The rostrum of the skull was relatively short compared to other pholidosaurids. Oceanosuchus was described in 2007 by Hua and colleagues. The type species is O. boecensis.

Pholidosaurus

Pholidosaurus is an extinct genus of neosuchian crocodylomorph. It is the type genus of the family Pholidosauridae. Fossils have been found in northwestern Germany. The genus is known to have existed during the Berriasian stage of the Early Cretaceous. Fossil material found from the Annero and Jydegård Formations in Skåne, Sweden and on the island of Bornholm, Denmark, have been referred to as a mesoeucrocodylian, and possibly represent the genus Pholidosaurus.

Sarcosuchus

Sarcosuchus (; meaning "flesh crocodile") is an extinct genus of crocodyliform and distant relative of living crocodylians that lived 112 million years ago. It dates from the early Cretaceous Period of what is now Africa and South America and is one of the largest crocodile-like reptiles that ever lived. It was almost twice as long as the modern saltwater crocodile and weighed up to 8 tonnes (7.9 long tons; 8.8 short tons).

The first remains were discovered during several expeditions led by the French paleontologist Albert-Félix de Lapparent, spanning from 1946 to 1959, in the Sahara. These remains were fragments of the skull, vertebrae, teeth, and scutes. In 1964, an almost complete skull was found in Niger by the French CEA, but it was not until 1997 and 2000 that most of its anatomy became known to science, when an expedition led by the American paleontologist Paul Sereno discovered six new specimens, including one with about half the skeleton intact and most of the spine.

Symptosuchus

Symptosuchus is an extinct genus of goniopholidid mesoeucrocodylian. It is known from the Late Cretaceous of Argentina. Argentine paleontologist Florentino Ameghino named the genus in 1899, along with the type species S. contortidens. It was formally described by Carlos Rusconi in 1934.

Terminonaris

Terminonaris is a genus of extinct pholidosaurid crocodyliforms that lived in the Late Cretaceous (Cenomanian and Turonian). The name means: “enlarged snout or nose” at the front of the skull. Terminonaris is an early crocodile, within a subgroup called Mesoeucrocodylia. Its remains have only been found in North America and Europe. Originally known under the generic name Teleorhinus, it was once believed to be a teleosaurid (a family of marine gavial-like thalattosuchians). Both, prehistoric crocodiles such as Terminonaris as well as modern crocodiles belong to the same group called crocodyliformes, although modern crocodiles have specific features that show indicate distant relatives of this species and in the subgroup Eusuchia.

Tethysuchia

Tethysuchia is an extinct clade of neosuchian mesoeucrocodylian crocodylomorphs from the late Middle Jurassic (Bathonian stage) to the Early Eocene (Ypresian stage) of Asia, Europe, North America and South America. It was named by the French paleontologist Eric Buffetaut in 1982 as a suborder. Tethysuchia was considered to be a synonym of Dyrosauridae or Pholidosauridae for many years. In most phylogenetic analyses the node Dyrosauridae+Pholidosauridae was strongly supported. De Andrade et al. (2011) suggested that Tethysuchia be resurrected for that node. They defined it as a node-based taxon "composed of Pholidosaurus purbeckensis (Mansel-Pleydell, 1888) and Dyrosaurus phosphaticus (Thomas, 1893), their common ancestor and all its descendants". In their analysis they found that the support for Tethysuchia is actually stronger than the support for Thalattosuchia. The following cladogram shows the position of Tethysuchia among the Neosuchia sensu this study.

Vectisuchus

Vectisuchus is a genus of goniopholidid mesoeucrocodylian, known from the Early Cretaceous-age Wealden Group of the Isle of Wight, England. It was a small, piscivorous crocodylomorph with a narrow, elongate snout, and relatively long forearms. The type specimen, SMNS 50984, was found in 1977. When discovered, it was complete and right-side-up, but the posterior portion was lost during excavation. Vectisuchus was described in 1980. The type species is V. leptognathus.

Wahasuchus

Wahasuchus is a genus of extinct mesoeucrocodylian of the Middle Campanian age found in the Quseir Formation, Egypt. The generic name derives from the Arabic word واحة (waha), which means "oasis", and souchos from the Greek in honor of crocodile-headed god of ancient Egypt. The specific egyptensis (Lat.) means from Egypt.Fossils of skull and jaw fragments, dorsal vertebrae, and fragmentary appendicular remains have been recovered.

Major groups within Mesoeucrocodylia
Notosuchia
Sebecia
Neosuchia
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