Enalioetes Temporal range: Early Cretaceous | |
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Skull of Enalioetes schroederi | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauria |
Clade: | Pseudosuchia |
Clade: | Crocodylomorpha |
Clade: | Crocodyliformes |
Suborder: | † Thalattosuchia |
Family: | † Metriorhynchidae |
Subfamily: | † Metriorhynchinae |
Genus: | † Enalioetes Sachs et al., 2024 |
Type species | |
†Enalioetes schroederi Sachs et al., 2024 | |
Synonyms | |
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Enalioetes is an extinct genus of metriorhynchid thalattosuchian from the Early Cretaceous Stadthagen Formation of Germany. The type species is E. schroederi. [1]
Prior to its description in 2024, it was known under the informal names Enaliosuchus "schroederi" and Cricosaurus "schroederi".
The holotype, which consists of much of the skull as well as parts of the neck, was discovered no later than 1916 in the Stadthagen Formation in Engelbostel, Germany. At the time of its discovery, the fossil material was thought to have belonged to a type of ichthyosaur by its discoverer, a government architect by the name of D. Hapke. Hapke then turned the fossils over to Prussian paleontologist Henry Schroeder in 1916, who proceeded to prepare and describe the material. Some years prior, in 1883, Ernst Koken described another metriorhynchid from the Stadthagen Formation, which he named Enaliosuchus macrospondylus . [2] Noting some similarities in the anatomy of the first two neck vertebrae, Schroeder placed the so-called "Sachsenhagen specimen" in the genus Enaliosuchus, but notably did not explicitly regard it as being the same species as S. macrospondylus nor declared it a distinct species instead. In his work Schroeder also briefly contemplated the possibility that Enaliosuchus was synonymous with the French metriorhynchid Neustosaurus , although he ultimately forewent any definitive conclusions due to the lack of overlapping fossil material. [3] [4]
Schroeder eventually returned the fossils to Mr. Hapke, though little is known on the whereabouts of the material during this period. Oskar Kuhn for instance erronously claimed that the material was housed in the capital of Berlin in his 1936 paper on the animal, when it had in fact been donated to the Mindener Museum in North Rhine-Westphalia. Regardless, Kuhn too contemplated the potential synonymity of Enaliosuchus and Neustosaurus, but did not yet lump the two genera. He did however recognize the "Sachsenhagen specimen" as a distinct form, giving it the name "Enaliosuchus schröderi". Kuhn did however not provide a proper diagnosis, instead simply referring readers to Schroeder's 1923 paper, [5] which only found minor differences that at the time were chalked up to changes that occurred during ontogeny. This actually represented a violation of the International Code of Zoological Nomenclature (ICZN), which states that taxa published after 1930 either require a direct, textual description of the diagnostic features or at least a bibliographic reference to such, rendering it a nomen nudum. Furthermore, Kuhn's name was a hypercorrection, using an Umlaut despite the fact that Schroeder's name is spelled with an "oe" rather than an "ö". Even if it wasn't, the use of diacritic marks is not considered acceptable by the ICZN, meaning that subsequent publications correctly spelled the name as "E. schroederi". [4]
While some authors continued to use the name E. schroederi, [6] some later research questioned whether or not the animal was distinct to begin with. First among those was a 1961 publication by Sickenberg, which cast doubt over the validity of "E. schroederi" due to Schroeder's own uncertainty regarding the "Sachsenhagen specimen"s affinities. In 2000 Hua and colleagues formally synonymized E. macrospondylus with "E. schroederi" [7] and in 2006 Karl and colleagues sunk all Enaliosuchus material into Metriorhynchus as indeterminate species. Contrary to this, Young and Andrade placed "E. schroederi" in the genus Cricosaurus as C. schroederi. [1] [4]
Eventually, first hand examination of the fossil material by Sachs et al. showed that both E. macrospondylus and "E. schroederi" were distinct species, separated primarily by the anatomy of the earliest neck vertebrae. The team further argued for a closer relationship between E. macrospondylus and Neustosaurus gigondarum, with the two animals possibly being congeneric, eventhough both taxa are for now considered nomina dubia. [8] "E. schroederi" was redescribed in 2024 by Sachs and colleagues, for the first time giving it an actual diagnosis and erecting the new genus Enalioetes. [4]
The genus name derives from the Greek "enalios" meaning "from the sea" and the suffix "-etes" to mean "dweller", a reference to the marine habits of metriorhynchids. The species name "schroederi" honours the initial work conducted on the animal by Henry Schroeder and is carried over from Kuhn's nomina nuda "Enaliosuchus schroederi". Sachs and colleagues argue that this avoids confusion in future studies, as the species name has a long history of use in scientific literature. [4]
The genus is among the most recent metriorhynchids known and one of the best preserved Cretaceous members of its group, preserving much of the skull as well as parts of the neck. [4]
Multiple phylogenetic analysis were run under different conditions, both using equal and implied weighting of the characters. Several of the analysis were run with the constraint of forcing Enalioetes into one of several groups (specifically the genus Cricosaurus, a clade of unnamed rhacheosaurins and the group formed by Enaliosuchus & Neustosaurus), which were in turn compared to the results yielded by conducting the same analysis without such constraints. The results of this method, known as the templeton test, show that it would take three additional steps relative to the unconstrained result to force Enalioetes into Cricosaurus, four to force it into the unnamed rhacheosaurin clade and only one for it to clade with Enaliosuchus and Neustosaurus. [4]
The results of the unconstrained analysis are shown in the tree below, which recovers Enalioetes deep within Metriorhynchidae. The tree recovers the family split into two major groups, Geosaurinae and Metriorhynchinae, with Enalioetes siting in a relatively basal position in the latter. It was recovered as more derived than Enaliosuchus and Neustosaurus but more basal than any rhacheosaurin group or Cricosaurus, the three clades it might also belong to. Sachs and colleagues note that the alternate placements could however not be excluded either. [4]
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There are multiple reasons for the ambiguity surrounding the placement of Enalioetes among metriorhynchids. For instance, few members of the clade are known from 3 dimensionally preserved skulls, making it uncertain how some internal skull characters were distributed (though they generally appear absent in Jurassic taxa). The lack of overlapping material with the other Cretaceous form is another hinderance, as is the lack of a cranial rostrum which was used as a key region for diagnostic features in the work of Young et al. (2024). Overall this limited comparative material means that the phylogenetic position of the animal is far from settled. [4]
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 yet. The name Dakosaurus means "biter lizard", and is derived from the Greek dakos ("biter") and σαῦρος -sauros ("lizard").
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.
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- and σαῦρος -sauros ("lizard"). The name Geosaurus was established by the French naturalist Georges Cuvier in 1824.
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 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.
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.
Emanuel Bunzel (1828–1895), was an Austrian paleontologist.
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.
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.
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.
Zoneait is an extinct genus of thalattosuchian crocodylomorph known from a single species, Zoneait nargorum, from the Middle Jurassic Snowshoe Formation of Oregon.
Eichstaettisaurus is a genus of lizards from the Late Jurassic and Early Cretaceous of Germany, Spain, and Italy. With a flattened head, forward-oriented and partially symmetrical feet, and tall claws, Eichstaettisaurus bore many adaptations to a climbing lifestyle approaching those of geckoes. The type species, E. schroederi, is among the oldest and most complete members of the Squamata, being known by one specimen originating from the Tithonian-aged Solnhofen Limestone of Germany. A second species, E. gouldi, was described from another skeleton found in the Matese Mountains of Italy. Despite being very similar to E. schroederi, it lived much later, during the Albian stage. Fossils of both species show exceptional preservation due to deposition in low-oxygen marine environments.
The Stadthagen Formation is a geological formation located in Hanover, Germany, dating to the Early Cretaceous (Valanginian). The formation contains fossils of reptiles and fish, suggesting that the formation was once a shallow marine setting.
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