Geosaurus | |
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Skull of G. giganteus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Pseudosuchia |
Clade: | Crocodylomorpha |
Suborder: | † Thalattosuchia |
Family: | † Metriorhynchidae |
Subtribe: | † Geosaurina |
Genus: | † Geosaurus Cuvier, 1824 |
Type species | |
†Geosaurus giganteus Sömmerring, 1816 | |
Species | |
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Synonyms | |
Geosaurus is an extinct genus of marine crocodyliform within the family Metriorhynchidae, that lived during the Late Jurassic and the Early Cretaceous. Geosaurus was a carnivore that spent much, if not all, its life out at sea. No Geosaurus eggs or nests have been discovered, so little is known of the reptile's lifecycle, unlike other large marine reptiles of the Mesozoic, such as plesiosaurs or ichthyosaurs which are known to give birth to live young out at sea. Where Geosaurus mated, whether on land or at sea, is currently unknown. The name Geosaurus means "Mother of Giants lizard", and is derived from the Greek Ge - ("Earth", the mythical mother of the Giants) and σαῦρος -sauros ("lizard"). The name Geosaurus was established by the French naturalist Georges Cuvier in 1824. [3]
Geosaurus was a large marine reptile of the group Crurotarsi, the line leading to modern crocodilians. It was about 2.5–3 m (8.2–9.8 ft) long. [4] Specifically, it was a "marine crocodile", or thalattosuchian. Geosaurus was similar in appearance to the related Dakosaurus with a relatively short skull and curved teeth designed for slashing, which it likely used to attack large prey.
Many early depictions of Geosaurus were based on a nearly complete specimen described by Eberhard Fraas, which Fraas classified as a distinct species of Geosaurus, G. suevicus. This specimen was found in Germany and dated to the late Jurassic (Late Kimmeridgian) period. [5] G. suevicus had a distinctively long, narrow snout filled with small, pointed teeth very different from skulls belonging to the type species. Further study and a redescription of Geosaurus published in 2009 showed that these long-snouted forms actually represent individuals of Cricosaurus . [6]
Genera considered junior synonyms of Geosaurus include Brachytaenius and Halilimnosaurus . [7] Numerous species had been assigned to this genus since the 19th Century. However, phylogenetic analyses begun in 2005 did not support the monophyly of Geosaurus. [8] [9] Although some traditional species, such as G. suevicus and G. araucanensis formed a natural group, [10] Enaliosuchus is also within that group. [8] [11] This, as well as further study showing that traditional metriorhynchid genera were not grouped based on actual relationships, necessitated almost all traditional species being removed from Geosaurus and reclassified elsewhere, as well as several species previously placed in other genera to be reclassified as species of Geosaurus. [6]
The species included below follow this revised classification, presented by Young and Andrade in 2009. [6]
A large species of Geosaurus is known only from a single tooth from the Nusplingen Plattenkalk of Germany. [14]
Cladogram after Cau & Fanti (2010). [15]
Geosaurus |
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In 2009, Young and de Andrade published a re-description of Geosaurus, examining its relationships and the validity of species lumped into the genus. After performing a phylogenetic analysis of metriorhynchids, they found that many species were grouped in a paraphyletic manner or with the wrong genera. Specifically, they found that several species formerly classified as Geosaurus, including G. suevicus, G. saltillense, [16] G. vignaudi, [17] and G. araucanensis [18] were actually examples of the related Cricosaurus . [6] Rhacheosaurus gracilis , another long-snouted species, was also at one time considered a species of Geosaurus (as G. gracilis). [6] [19]
Geosaurus carpenteri, also formerly referred to Dakosaurus, is known from a partial skull. Teeth identical to the known teeth of this species are also known from Yorkshire, UK. It was assigned to its own genus, Torvoneustes by Andrade et al., 2010. [14]
An unnamed specimen classified as Geosaurus was found in the Oxfordian Jagua Formation of Cuba, [20] though further study has shown this species to be more closely related to Cricosaurus as well. [6]
Additionally, Geosaurus fossils have been reported from the Vaca Muerta of Argentina. [21]
Several species of metriorhynchids are known from the Mörnsheim Formation (Solnhofen limestone, early Tithonian) of Bavaria, Germany: Geosaurus giganteus, Dakosaurus maximus , Cricosaurus suevicus and Rhacheosaurus gracilis . It has been hypothesised that niche partitioning enabled several species of crocodyliforms to co-exist. The top predators of this Formation appear to be G. giganteus and D. maximus, which were large, short-snouted species with serrated teeth. The long-snouted C. suevicus and R. gracilis would have fed mostly on fish, although the more lightly built Rhacheosaurus may have specialised towards feeding on small prey. In addition to these four species of metriorhynchids, a moderate-sized species of Steneosaurus was also contemporaneous. [22]
Dakosaurus is an extinct genus of crocodylomorph within the family Metriorhynchidae that lived during the Late Jurassic and Early Cretaceous. It was large, with teeth that were serrated and compressed lateromedially. The genus was established by Friedrich August von Quenstedt in 1856 for an isolated tooth named Geosaurus maximus by Theodor Plieninger in 1846. Dakosaurus was a carnivore that spent much, if not all, its life out at sea. The extent of its adaptation to a marine lifestyle means that it is most likely that it mated at sea, but since no eggs or nests have been discovered that have been referred to Dakosaurus, whether it gave birth to live young at sea like dolphins and ichthyosaurs or came ashore like turtles is not known. The name Dakosaurus means "biter lizard", and is derived from the Greek dakos ("biter") and σαῦρος -sauros ("lizard").
Aggiosaurus is an extinct genus of geosaurine metriorhynchid crocodyliform known from the Late Jurassic of Nice, southeastern France. It contains a single species, Aggiosaurus nicaeensis, which was named by H. Ambayrac in 1913.
Machimosaurus is an extinct genus of machimosaurid crocodyliform from the Late Jurassic and Early Cretaceous. The type species, Machimosaurus hugii, was found in Switzerland. Other fossils have been found in England, France, Germany, Portugal, Switzerland and Tunisia. Machimosaurus rex is the largest named teleosauroid and thalattosuchian, with an estimated length of up to 7.15 m (23.5 ft). Machimosaurus is the largest known crocodyliform of the Jurassic.
Metriorhynchus is an extinct genus of marine crocodyliform that lived in the oceans during the Late Jurassic. The type species, M. brevirostris was named in 1829 as a species of Steneosaurus before being named as a separate genus by the German palaeontologist Christian von Meyer in 1832. The name Metriorhynchus means "Moderate snout", and is derived from the Greek Metrio- ("moderate") and -rhynchos ("snout").
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.
Enaliosuchus is a dubious genus of extinct marine crocodyliform within the family Metriorhynchidae that lived during the Valanginian stage of the Early Cretaceous. It is known from fossil remains found in France and Germany and it was first described in 1883,. The name Enaliosuchus means "Marine crocodile", and is derived from the Greek Enalios- ("marine") and -suchos ("crocodile"). Two species are known: the type species E. macrospondylus, described in 1883, and the second species E. schroederi, described in 1936, which was likely the same animal as E. macrospondylus.
Teleidosaurus is an extinct genus of carnivorous metriorhynchoid crocodyliform from Middle Jurassic deposits of Normandy, France. The name Teleidosaurus means "Complete lizard", and is derived from the Greek Teleidos- ("complete") and σαῦρος -sauros ("lizard").
Thalattosuchia is a clade of 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.
Neustosaurus is a genus of marine crocodyliform from the Early Cretaceous. Neustosaurus was a carnivore that spent much, if not all, its life out at sea. No Neustosaurus eggs or nest have been discovered, so little is known of the reptile's lifecycle, unlike other large marine reptiles of the Mesozoic, such as plesiosaurs or ichthyosaurs which are known to give birth to live young out at sea. Where Neustosaurus mated, whether on land or at sea, is currently unknown. The name Neustosaurus means "Swimming lizard", and is derived from the Greek Neustos- ("swimming") and σαῦρος -sauros ("lizard").
Purranisaurus is an extinct genus of marine crocodyliform from the Middle to Late Jurassic period of Chile and Vaca Muerta of Argentina. Rusconi originally regarded Purranisaurus potens to be a plesiosaur; however, Gasparini demonstrated that it was in fact a metriorhynchid crocodyliform, and that may be a junior synonym of Metriorhynchus. It was about 3.3 m (11 ft) long.
Plesiosuchus is an extinct genus of geosaurine metriorhynchid crocodyliform known from the Late Jurassic of Dorset, England and possibly also Spain. It contains a single species, Plesiosuchus manselii.
Rhacheosaurus is an extinct genus of marine crocodyliform belonging to the family Metriorhynchidae. The genus was established by Christian Erich Hermann von Meyer in 1831 for skeletal remains from the Tithonian of Germany. It was a relatively small reptile, measuring between 1.39 and 1.57 m long.
Suchodus is an extinct genus of marine crocodyliform from the Middle to Late Jurassic period of England and France. It measured between 3.3 and 4.7 m in total body length.
Cricosaurus is an extinct genus of marine crocodyliforms of the Late Jurassic. belonging to the family Metriorhynchidae. The genus was established by Johann Andreas Wagner in 1858 for three skulls from the Tithonian of Germany. The name Cricosaurus means "Ring lizard", and is derived from the Greek Krikos- ("ring") and σαῦρος -sauros ("lizard"). It was a relatively small reptile, with C. suevicus and C. araucanensis measuring 2 m (6.6 ft) and 3.2 m (10 ft) in total body length, respectively.
Aeolodon is an extinct genus of teleosaurid crocodyliform reptile from the Late Jurassic (Tithonian) of Germany and France that was initially named as a species of Crocodylus in 1814. Although previously synonymized with Steneosaurus, recent cladistic analysis considers it distantly related to the Steneosaurus type species and the type species is A. priscus, named in 1830 and described in 2020.
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.
Tyrannoneustes is an extinct genus of geosaurine metriorhynchid crocodyliform from the Callovian stage Oxford Clay Formation of England and the Marnes de Dives of France. It contains a single species, Tyrannoneustes lythrodectikos, meaning "blood-biting tyrant swimmer".