Goniopholididae Temporal range: Calsoyasuchus is a goniopholidid Possible Early Jurassic record if | |
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Amphicotylus milesi from the Late Jurassic of North America | |
Skull of Goniopholis | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Crocodylomorpha |
Clade: | Crocodyliformes |
Clade: | Neosuchia |
Family: | † Goniopholididae Cope, 1875 |
Subgroups | |
Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle Jurassic (possibly Early Jurassic, see below) and the Late Cretaceous.
The dorsal armour of goniopholidids is composed of two rows of paired osteoderms (as opposed to the four main pairs present in living crocodilians and other eusuchians), which are rectangular in shape and wider than they are long, with the lateral margins ventrally deflected and an anterior process for a ‘peg and groove’ articulation. [1] Unlike modern crocodilians they have ventral osteoderms as well. Their forelimbs are also proportionally very long, particularly in the humeri and wrist bones, being as long or longer than the hindlimbs, the opposite of the condition seen in modern crocodilians. Some like Anteophthalmosuchus also have forwardly oriented eyes, as opposed to the dorsally oriented eyes seen in modern forms. These suggest multiple biomechanical differences from modern species. [2] [3]
Goniopholidids likely had a similar ecology to modern crocodilians as semi-aquatic ambush predators. [4]
Goniopholidids have only been found in Laurasia (Asia, Europe and North America). The oldest possible member of the group is Calsoyasuchus from the Early Jurassic of North America. [5] However, its placement is disputed, with some studies recovering it as only distantly related to goniopholidids. [6] [7] The goniopholidids were present across Eurasia during the Middle Jurassic [8] [9] and were widespread in North America during the Late Jurassic [10] and continued to remain prominent across Laurasia during the Early Cretaceous. [4] [11] Goniopolidids persisted into the late Upper Cretaceous in North America based on Denazinosuchus, from the Campanian-Maastrichtian of New Mexico, which is only known from fragmentary remains, and has been disputed as a member of the group, as well as remains of an unnamed goniopholidid from the Campanian aged Aguja Formation of Texas. [12]
The following cladogram simplified after an analysis presented by Marco Brandalise de Andrade and colleagues in 2011. [13]
Neosuchia |
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Saltoposuchus is an extinct genus of small, long-tailed crocodylomorph reptile (Sphenosuchia), from the Norian of Europe. The name translated means "leaping foot crocodile". It has been proposed that Terrestrisuchus gracilis and Saltoposuchus connectens represent different ontogenetic stages of the same genus. Saltoposuchus was commonly referred to in popular literature as the ancestor to dinosaurs; however, recent scientific research shows that this is not the case.
A dermal bone or investing bone or membrane bone is a bony structure derived from intramembranous ossification forming components of the vertebrate skeleton, including much of the skull, jaws, gill covers, shoulder girdle, fin rays (lepidotrichia), and the shells of turtles and armadillos. In contrast to endochondral bone, dermal bone does not form from cartilage that then calcifies, and it is often ornamented. Dermal bone is formed within the dermis and grows by accretion only – the outer portion of the bone is deposited by osteoblasts.
Goniopholis is an extinct genus of goniopholidid crocodyliform that lived in Europe and North America during the Late Jurassic and Early Cretaceous. Like other goniopholidids, it resembled living crocodilians, and probably had a similar ecology as semi-aquatic ambush predators.
Metriorhynchidae is an extinct family of specialized, aquatic metriorhynchoid crocodyliforms from the Middle Jurassic to the Early Cretaceous period of Europe, North America and South America. The name Metriorhynchidae was coined by the Austrian zoologist Leopold Fitzinger in 1843. The group contains two subfamilies, the Metriorhynchinae and the Geosaurinae. They represent the most marine adapted of all archosaurs.
Thalattosuchia is a clade of mostly marine crocodylomorphs from the Early Jurassic to the Early Cretaceous that had a cosmopolitan distribution. They are colloquially referred to as marine crocodiles or sea crocodiles, though they are not members of Crocodilia and records from Thailand and China suggest that some members lived in freshwater. The clade contains two major subgroupings, the Teleosauroidea and Metriorhynchoidea. Teleosauroids are not greatly specialised for oceanic life, with back osteoderms similar to other crocodyliformes. Within Metriorhynchoidea, the Metriorhynchidae displayed extreme adaptions for life in the open ocean, including the transformation of limbs into flippers, the development of a tail fluke, and smooth, scaleless skin, and probably gave live birth, seemingly uniquely among archosaurs.
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 than 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.
Calsoyasuchus is a genus of crocodylomorph that lived in the Early Jurassic. Its fossilized remains were found in the Sinemurian-Pliensbachian-age Kayenta Formation on Navajo Nation land in Coconino County, Arizona, United States. Formally described as C. valliceps, it is known from a single incomplete skull which is unusually derived for such an early crocodile relative. This genus was described in 2002 by Ronald Tykoski and colleagues; the specific name means "valley head" and refers to a deep groove along the midline of the nasal bones and frontal bones. It has often been interpreted as the earliest diverging member of Goniopholididae, but other studies have recovered it in various other positions.
The Adamantina Formation is a geological formation in the Bauru Basin of western São Paulo state, in southeastern Brazil.
Eutretauranosuchus is an extinct genus of goniopholidid crocodyliform. E. delfsi is the only known species within the genus.
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 than to Notosuchus terrestris. Members of Neosuchia generally share a crocodilian-like bodyform adapted to freshwater aquatic life, as opposed to the terrestrial habits of more basal crocodylomorph groups. The earliest neosuchian is suggested to be the Early Jurassic Calsoyasuchus, which lived during the Sinemurian and Pliensbachian stages in North America. It is often identified as a member of Goniopholididae, though this is disputed, and the taxon may lie outside Neosuchia, which places the earliest records of the group in the Middle Jurassic.
Amphicotylus is an extinct genus of goniopholidid mesoeucrocodylian from the Tithonian of Colorado, Wyoming, and Oklahoma. It was described in 1878.
Sunosuchus is an extinct genus of goniopholidid mesoeucrocodylian. Fossils are known from China, Kyrgyzstan, and Thailand and are Jurassic in age, although some may be Early Cretaceous. Four species are currently assigned to the genus: the type species S. miaoi and the species S. junggarensis, S. shartegensis, and S. shunanensis. All species are from China. Goniopholis phuwiangensis, also from Thailand, was reassigned to Sunosuchus by Andrade et al. (2011). The material from Kyrgyzstan has not been assigned to any species.
Metriorhynchoidea is an extinct superfamily of thalattosuchian crocodyliforms from the Early Jurassic to the Early Cretaceous of Europe, North America and South America. Metriorhynchids are fully aquatic crocodyliforms. Named by Fitzinger, in 1843, it contains the basal taxa like Teleidosaurus, Zoneait and Eoneustes and the family Metriorhynchidae. An unnamed taxon is known from Chile.
Torvoneustes is an extinct genus of metriorhynchid thalattosuchian. It is known from skull and postcranial remains found in the Kimmeridge Clay Formation of Dorset and Wiltshire, England, the Virgula Marls of Switzerland and also from Oaxaca, Mexico . The holotype skull of the type species was initially assigned to the species Metriorhynchus superciliosus. Postcranial remains were later discovered from the same quarry as the skull, and then these specimens were recognised as belonging to a new species of Dakosaurus, as D. carpenteri. The species was named to honour Simon Carpenter, an amateur geologist from Frome in Somerset, who discovered the fossils.
Geosaurinae is a subfamily of metriorhynchid crocodyliforms from the Middle Jurassic to the Early Cretaceous of Europe, North America and South America. Named by Richard Lydekker, in 1889, it contains the metriorhynchids Suchodus, Purranisaurus, Neptunidraco, Tyrannoneustes, Torvoneustes, Dakosaurus, Geosaurus and Plesiosuchus. The last four taxa form a tribe within Geosaurinae, the Geosaurini. Geosaurinae is one of two subfamilies of Metriorhynchidae, the other being Metriorhynchinae.
Metriorhynchinae is a subfamily of metriorhynchid crocodyliforms from the late Middle Jurassic to the Early Cretaceous of Europe, North America and South America. Named by Fitzinger, in 1843, it contains the metriorhynchids Maledictosuchus, Gracilineustes, Metriorhynchus, Cricosaurus and Rhacheosaurus. The last three taxa form a tribe within Metriorhynchinae, the Rhacheosaurini. Metriorhynchinae is one of two subfamilies of Metriorhynchidae, the other being Geosaurinae.
Anteophthalmosuchus is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England, eastern Spain, and western Belgium.
Hulkepholis is an extinct genus of goniopholidid mesoeucrocodylian from the Early Cretaceous of southern England and eastern Spain. It contains two species, the type species, Hulkepholis willetti, and also H. plotos. Hulkepholis is most closely related to both species of Anteophthalmosuchus.
Chalawan is an extinct genus of pholidosaurid mesoeucrocodylian known from the Late Jurassic or Early Cretaceous Phu Kradung Formation of Nong Bua Lamphu Province, northeastern Thailand. It contains a single species, Chalawan thailandicus, with Chalawan shartegensis as a possible second species.
Coelognathosuchia is an extinct clade of neosuchian crocodyliforms that includes all taxa more closely related to the family Pholidosauridae than to Bernissartia fagesii or Eusuchia. 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.