This article relies largely or entirely on a single source .(January 2023) |
Turnersuchus Temporal range: Early Pliensbachian, | |
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Holotype of T. hingleyae on display at the Lyme Regis Museum | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Pseudosuchia |
Clade: | Crocodylomorpha |
Suborder: | † Thalattosuchia |
Genus: | † Turnersuchus Wilberg et al., 2023 |
Type species | |
†Turnersuchus hingleyae Wilberg et al., 2023 |
Turnersuchus is an extinct genus of thalattosuchian, a group of marine crocodylomorphs, from the Pliensbachian of the United Kingdom. It is the oldest diagnostic member of Thalattosuchia and was also found to be the group's most basal member, being situated outside the two major groups Metriorhynchoidea and Teleosauroidea. Subsequently, this genus is considered to be of great importance to understanding the relationship between thalattosuchians and other crocodylomorphs as well as their rapid diversification during the early Jurassic. Turnersuchus is a monotypic genus, meaning it includes only a single species, Turnersuchus hingleyae.
The fossils of Turnersuchus were discovered in 2017 within the Belemnite Marl Member of the British Charmouth Mudstone Formation in Dorset and the specimens were recorded under the West Dorset Fossil Collecting Code due to their rarity. The holotype, LYMPH 2021/45, is preserved in five large blocks with several additional isolated elements and consists of a partial skull and mandible, articulated cervical and dorsal vertebrae alongside various ribs of this region, isolated tail vertebrae, parts of the right shoulder girdle with the humerus and ulna, a tibia and a single osteoderm of the dorsal armor. [1] As of August 2023, the holotype is on display at the Lyme Regis Museum.
The name Turnersuchus hingleyae honors two people involved with the discovery of this taxon. The genus is named after Paul Turner who discovered the initial fossil blocks and subsequently donated them, while the species name is based on Elizabeth "Lizzie" Hingley who prepared the specimen and discovered additional material. The suffix -suchus is derived from the Greek "soukhos" for crocodile. [1]
Although the skull is primarily known from elements of the braincase, the fragmentary remains of the mandible show that Turnersuchus had rather narrow jaws. However, the authors note that the basioccipital tubera, which are typically associated with elongation of the jaws and longirostrine skull morphology, are only poorly developed. They subsequently suggest that although narrow, the jaws of Turnersuchus may not have been as long as in some of the more derived thalattosuchians. [1]
The shoulder girdle is only partially preserved. The shoulder blade is the better preserved element, being present in its entirety and only broken, while the coracoid is only known from its proximal end. Generally, these elements are larger in Turnersuchus than in some metriorhynchids such as Cricosaurus , which has reduced forelimbs. The preserved portion of the coracoid is similar to the teleosaur Charitomenosuchus and the metriorhynchid Magyarosuchus , while the flattened scapula bears resemblance to Macrospondylus and Pelagosaurus with the almost equally expanded distal and proximal ends. The humerus is flat and broad with a nearly straight shaft and reduced deltopectoral crest. The ulna is strongly curved and possesses a proximal surface twice as wide as the shaft of the bone. Compared to the humerus, the ulna is highly reduced, only half the length of the former. The reduction of the deltapectoral crest and the length of the ulna are also seen in later metriorhynchids, however to a much greater degree. [1]
The only known osteoderm of Turnersuchus is oval in shape with pitted ornamentation and a slight keel running down the middle. Based on its shape it is believed that it may have been a tail osteoderm closer to the end than the base, which are rarely reported from thalattosuchians. [1]
Two phylogenetic analysis were conducted to determine the position of Turnersuchus within Thalattosuchia, one based on the dataset of Wilberg et al. (2019) and another modified from Herrera et al. (2021). Both datasets result in Turnersuchus being recovered as the basalmost member of Thalattosuchia, supported by four and five synapomorphies respectively. While the inclusion of Turnersuchus within Thalattosuchia is thus well supported, the exclusion of it from either teleosauroids or metriorhynchoids is less so. Only a single synapomorphy unambiguously points towards Turnersuchus not being part of these clades in either analysis. These traits additionally differ in both datasets, with Wilberg et al. excluding it based on a character of the basioccipital and Herrera et al. excluding it based on the shape of the scapula. Each analysis thus offers three alternative placements within Thalattosuchia, all of which being only one step longer than the most parsimonious position. These include the possibility that Turnersuchus may be the earliest diverging metriorhynchoid, earliest diverging teleosauroid or most closely allied with Plagiophthalmosuchus . The last placement differs in its details between the analysis. In the Wilberg dataset this alternative would place both taxa at the base of teleosauroids, while the Herrera dataset suggests that in this alternative placement Plagiopthalmosuchus may be an even more basal thalattosuchian than Turnersuchus. Overall, while both analysis agree that the position as basalmost member of Thalattosuchia is the most parsimonious result, this may change as more species are described. [1]
The most parsimonious trees of either analysis are shown below. On the left the strict consensus tree based on the Wilberg et al. dataset and on the right the strict consensus tree of the Herrera et al. dataset.
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The discovery of Turnersuchus helps fill two major gaps in the understanding of Thalattosuchia. Previously, little was known on their origin, diversification and their relationship to other groups due to the lack of basal members or other transitional forms linking them to Crocodylomorpha as a whole. Phylogenetic analysis often yield contradictory results primarily nesting them in three different positions. As a sister to Crocodyliformes (as also recovered for Turnersuchus using the Wilberg dataset), as more derived mesoeucrocodylians (similar to their position in the Herrera dataset) or as derived neosuchians allied with pholidosaurids and dyrosaurids. The understanding of their dispersal is also heavily affected by the lacking fossil record of early thalattosuchians. Prior to Turnersuchus, the oldest known thalattosuchians were Toarcian in age, at which point the group was already split into its two main branches, rich in species and found across several continents. [1]
With Turnersuchus being recovered as the basalmost thalattosuchian, several similarities were recognized between it and basal members of both Teleosauroidea and Metriorhynchoidea, namely Plagiophthalmosuchus and Pelagosaurus. This helps solidify these traits as ancestral (plesiomorphic) traits of the group as a whole. These ancestral traits include the presence of large supratemporal fenestrae, unflattened skull table, temporal bars with ornamented surface, a squamosal facet, fused pterygoids and the prootic being broadly exposed along the side of the braincase. The quadrate in particular shows several ancestral traits between the three taxa. It makes up the front, top and lower margin of the external otic aperture, excluding the squamosal bone from contributing to its margins and it is furthermore overlapped by a broad process of the otoccipital bone. The quadrate however also serves to differentiate Turnersuchus from more derived thalattosuchians, as it is less integrated into the braincase. This is of significance due to the general evolutionary trend of crocodylomorphs progressively developing quadrates that articulate more and more with the bones of the braincase. This trend can be observed throughout the paraphyletic sphenosuchians and is later continued by crocodyliforms. Thalattosuchians show an intermediate condition between the two, with Turnersuchus displaying a quadrate that is clearly less well integrated than in crocodyliforms or even more derived thalattosuchians. The development of the basioccipital tubera may be ancestral trait that Turnersuchus displays. These tubera are generally thought to be associated with the development of elongated jaws, the longirostrine condition, and are not unique to thalattosuchians. They are however consistently found in members of the clade due to their longirostrine skulls, the only exceptions being the short-skulled Dakosaurus andiniensis and Turnersuchus. Although Turnersuchus had slender jaws, the poorly developed tubera indicate that they were not as elongated as in later forms and that thalattosuchians were ancestrally short-snouted. [1]
In addition to helping uncover the ancestral morphology of Thalattosuchia, Turnersuchus further aids in filling the ghost lineage leading up to the groups explosive radiation during the late Early Jurassic. It pushes the known record of thalattosuchians from the Toarcian back into the Pliensbachian, shortening the period of time between the group's first estimated appearance and the first confirmed fossils. However, the precise divergence remains uncertain. Bayesian analysis based on the two phylogenetic analysis yield vastly different results. Following the interpretation that thalattosuchians are early diverging mesoeucrocodylians, as in the Herrera dataset, the remaining ghost lineage would be relatively short, extending only into the Sinemurian stage of the early Jurassic. However, if they are indeed a sister group to crocodyliforms, as indicated by the Wilberg dataset, then the origin of Thalattosuchia would be pushed back into the Norian, over 13 million years earlier than indicated by the other analysis. The maximum ranges (95% highest posterior density) for both analysis would suggest that the group could have originated in the Late Triassic, either during the Norian or the Rhaetian. This is broadly confirmed by an in-press paper describing an indeterminate teleosauroid from the earliest Jurassic (Hettangian to Sinemurian) of Morocco, which would support the presence of thalattosuchians prior to the Pliensbachian with a possible origin prior to the beginning of the Jurassic. [1]
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").
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.
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.
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.
Junggarsuchus is an extinct genus of sphenosuchian crocodylomorph from the Middle or Late Jurassic period of China. The type and only species is J. sloani. The generic name of Junggarsuchus comes from the Junggar Basin, where the fossil was found, and the Greek word "souchos" meaning crocodile. The specific name, "sloani" is in honor of C. Sloan, who is credited with finding the holotype.
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.
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.
Steneosaurus is a dubious genus of teleosaurid crocodyliform from the Middle or Late Jurassic of France. The genus has been used as a wastebasket taxon for thalattosuchian fossils for over two centuries, and almost all known historical species of teleosauroid have been included within it at one point. The genus has remained a wastebasket, with numerous species still included under the label ‘Steneosaurus’, many of which are unrelated to each other.
Chacaicosaurus is a genus of neoichthyosaurian ichthyosaur known from the Middle Jurassic of Argentina. The single known specimen of this genus was excavated from the Los Molles Formation in Neuquén Province, and is housed at the Museo Olsacher under the specimen number MOZ 5803. This specimen consists of a skull, forelimb, some vertebrae, and some additional postcranial elements. The genus was named by Marta Fernández in 1994, and contains a single species, Chacaicosaurus cayi, making it the first named distinctive ichthyosaur from the Bajocian stage. It is a medium-sized ichthyosaur with a very long snout, which bears a ridge running along each side. The forelimbs of Chacaicosaurus are small and contain four main digits.
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.
Macrospondylus is an extinct genus of machimosaurid teleosauroid crocodyliform from the Early Jurassic (Toarcian) of Europe. Fossils are known from the Posidonia Shale of Germany, the Whitby Mudstone of the United Kingdom, and the "schistes bitumineux" of Luxembourg.
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.
Bathysuchus is an extinct genus of teleosaurid thalattosuchian from Late Jurassic (Kimmeridgian) deep water marine deposits in England and France. Bathysuchus displays features that suggest it was more pelagic than other teleosaurids, including smoother skull bones and reduced armour plating, similar to the fully marine metriorhynchids. This was possibly an adaptation to rising sea levels during the Kimmeridgian, as its earlier relatives such as Teleosaurus were suited for shallow coasts and lagoon environments.
Almadasuchus is an extinct genus of crocodylomorph known from the early Late Jurassic Puesto Almada Member of the e Cañadón Calcáreo Formation of Patagonia, Argentina. It contains a single species, Almadasuchus figarii. It is known from the holotype MPEF-PV 3838, a well-preserved posterior region of the skull as well as other skull and postcranial remains. Almadasuchus was recovered from Puesto Almada, 30 m above the fish beds, dated as Oxfordian in age.
Lemmysuchus is a genus of machimosaurid thalattosuchian from the Middle Jurassic Callovian of England and France. Like many other teleosauroids from Europe, it has had a convoluted taxonomic history.
Zoneait is an extinct genus of thalattosuchian crocodylomorph known from a single species, Zoneait nargorum, from the Middle Jurassic of Oregon. Z. nargorum was named in 2015 by paleontologist Eric Wilberg on the basis of several partial skulls, vertebrae, and forelimb bones that were found in an outcrop of the Snowshoe Formation near the town of Izee. It is a member of Metriorhynchoidea, a clade of marine-adapted thalattosuchians that existed until the Early Cretaceous. The skeleton of Zoneait possesses several adaptations for offshore marine life but retains features characteristic of its land-living ancestors, indicating that it is a transitional form between the fully marine metriorhynchids of the late Middle Jurassic to Early Cretaceous, and earlier non-marine crocodylomorphs. The Snowshoe Formation was deposited in a shallow marine environment within a tropical forearc basin, suggesting that Zoneait was a marine predator.
Indosinosuchus is a genus of teleosaurid neosuchian that lived during the Late Jurassic or Early Cretaceous in what is now Thailand. It contains two species, the type species I. potamosiamensis and I. kalasinensis, both recovered from the lower Phu Kradung Formation. It is unique among teleosauroids as it is the only named genus known from a freshwater environment, while most other members of the group are marine. Indosinosuchus is placed in the family Teleosauridae, but has a relatively robust skull that bears resemblance to members of the Machimosauridae. Biomechanical analysis of its mandible and teeth suggest that it would have had a substantial bite force comparable to animals like Lemmysuchus. The two Indosinosuchus species however differ in the speed at which they could open and close their jaws, impacting their respective ecology and possibly explaining how they coexisted in the same environment. All known specimens of this genus were recovered from a single locality, which has been interpreted as a mass death site, possibly caused by a drought or flash flood. The precise age of Indosinosuchus is unclear, as the vertebrate fossils of the Phu Kradung Formation support a Late Jurassic age, while palynological data suggests an Early Cretaceous (Berriasian) age.
Machimosauridae is an extinct family of teleosauroid thalattosuchian crocodyliforms. The family was first identified in 2016, when fossils of teleosauroid thalattosuchians, including an indeterminate close relative of Lemmysuchus and Machimosaurus, were described from the Middle Jurassic (Bathonian) of Morocco. The family was largely expanded in 2020 when the systematics of Teleosauroidea were re-reviewed. Members of this family generally were larger than the teleosaurids.