Sapheosaurids Temporal range: | |
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Skeleton of Piocormus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | Rhynchocephalia |
Suborder: | Sphenodontia |
Family: | † Sapheosauridae Baur 1895 |
Genera | |
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Sapheosaurs are an extinct group of rhynchocephalian reptiles from the Late Jurassic period. "Sapheosaurs" is an informal name for a group of rhynchocephalians closely related to the genus Sapheosaurus . It was first recognized as a group containing multiple genera by Hoffstetter in 1955. The group has sometimes been given a formal taxonomic name as the family Sapheosauridae, although in some analyses this group belongs to the family Sphenodontidae (which also contains the tuatara, the only living rhynchocephalian) and thus cannot be assigned its own family. They were fairly advanced rhynchocephalians which may have had semiaquatic habits. [1]
The most well-known members of the group are Sapheosaurus and Kallimodon , and most phylogenetic analyses on rhynchocephalians only study these two genera as representatives of sapheosaurs. Although a few early phylogenies in the 1990s did not find that these two formed a natural clade, [2] the relation between these two is now considered to be more stable, and has been found in practically every major analysis of rhynchocephalians since Apesteguía & Novas (2003). [3]
However, the relation between sapheosaurs as a whole and other rhynchocephalians is less clear. Although they are clearly members of the group Sphenodontia like almost all other rhynchocephalians, the construction of their jaw joints means that they were unable to move their jaws in a front-to-back chewing movement. This excludes them from the clade of sphenodonts which are capable of such movement, the so-called "eupropalinal sphenodonts" such as the tuatara, (formally known as Sphenodon), its close relatives, and the herbivorous opisthodonts. [1] They are also generally considered to be more derived (as in closer to Sphenodon) than clevosaurs and basal genera such as Godavarisaurus and Sphenocondor . The in-group relations of sphenodonts are inconsistent between analyses, so although sapheosaurs may be the sister group of eupropalinal sphenodonts under some methodologies, other potentially more derived taxon include Homoeosaurus , [4] Pamizinsaurus , Ankylosphenodon , the Sphenovipera + Theretairus clade, and pleurosaurs. [5]
A 2017 study utilizing both parsimony and bayesian analyses found some support for a clade including sapheosaurs, Vadasaurus , and pleurosaurs. Although the bayesian analysis placed this clade in a large polytomy with various other rhynchocephalian groups and genera, the parsimony analysis actually placed it among the eupropalinal sphenodonts. If this phylogeny is accurate, this would mean that an ancestor of this clade lost front-to-back chewing which was present in an even earlier ancestor, rather than never having it in the first place. [6] A 2022 study found a clade containing pleurosaurs, Vadasaurus, sapheosaurs,Homoeosaurus, Kallimodon and Leptosaurus, to the exclusion of other neosphenodontians like Sphenodontidae. This clade was called Leptorhynchia. [7]
All known members of the group lived in coastal environments of the late Jurassic in what is now Germany and France. They are very similar to each other, only noticeably differing in certain proportions of the skull. Evans (1988) and Ahmad (1993) have even considered that they may all belong to a single species, although a shortage of good studies and descriptions focusing on this group in particular means that any conclusions on the relationships between different sapheosaurs are uncertain at best. [1]
The postorbital region of the skull (behind the eyes) is lengthened and expanded in this group to such a point that it is longer than the preorbital region (in front of the eyes). Although clevosaurs also had skulls which were largest in the postorbital region, they evolved such a feature through different means, namely the preorbital region being shortened. [1] The upper temporal fenestrae (a pair of large holes on the top of the rear part of the skull) are long but fairly thin, a feature also known in Vadasaurus and Palaeopleurosaurus, potential relatives of the group. [6] Each upper temporal arch, which separates the upper temporal fenestrae from the lower temporal fenestrae (on the sides of the skull), is broad. Sapheosaurs, like other sphenodonts, had acrodont teeth which grew directly from the bone and could not be replaced. Historically Sapheosaurus was thought to be toothless. [1] However, it was later shown that Sapheosaurus and Oenosaurus possessed tooth plates on the upper and lower jaws, [8] formed by the fusion of maxillary and dentary teeth. Tooth plates are now thought to be characteristic of all sapheosaurs. [7] They are thought to have been used for crushing hard shelled organisms (durophagy). [9]
The centra (main body) of each caudal (tail) vertebra is flattened from the side in sapheosaurs. Sapheosaur vertebrae also had swollen neural arches (the area above the spinal cord) and zygapophyses (connecting joint plates), features also present in Ankylosphenodon. [1] Some have proposed that sapheosaurs were at least partially aquatic due to some similarities and/or possible close relations to Ankylosphenodon (now believed to be convergently evolved) or Vadasaurus and pleurosaurs. However, they do not share many of the adaptations that these other taxa possess, and some researchers are not convinced by this hypothesis. [10]
The Lepidosauria is a subclass or superorder of reptiles, containing the orders Squamata and Rhynchocephalia. Squamata includes lizards and snakes. Squamata contains over 9,000 species, making it by far the most species-rich and diverse order of non-avian reptiles in the present day. Rhynchocephalia was a formerly widespread and diverse group of reptiles in the Mesozoic Era. However, it is represented by only one living species: the tuatara, a superficially lizard-like reptile native to New Zealand.
Rhynchocephalia is an order of lizard-like reptiles that includes only one living species, the tuatara of New Zealand. Despite its current lack of diversity, during the Mesozoic rhynchocephalians were a speciose group with high morphological and ecological diversity. The oldest record of the group is dated to the Middle Triassic around 238 to 240 million years ago, and they had achieved a worldwide distribution by the Early Jurassic. Most rhynchocephalians belong to the group Sphenodontia ('wedge-teeth'). Their closest living relatives are lizards and snakes in the order Squamata, with the two orders being grouped together in the superorder Lepidosauria.
Sphenodontidae is a family within the reptile group Rhynchocephalia, comprising taxa most closely related to the living tuatara. Historically the taxa included within Sphenodontidae have varied greatly between analyses, and the group has lacked a formal definition. Cynosphenodon from the Jurassic of Mexico has consistently been recovered as a close relative of the tuatara in most analyses, with the clade containing the two and other very close relatives of the tuatara often called Sphenodontinae. The herbivorous Eilenodontinae, otherwise considered part of Opisthodontia, is considered to be part of this family in many recent studies as the sister group to Sphenodontinae. The earliest Sphenodontines are known from the Early Jurassic of North America, with other remains known from the Late Jurassic of Europe, the Late Cretaceous of South America and the Miocene-recent of New Zealand. Sphenodontines are characterised by a complete lower temporal bar caused by the fusion of a forward directed process (extension) of the quadrate/quadratojugal and the jugal, which was an adaptation for reducing stress in the skull during hard biting. Other synapomorphies of Sphenodontinae include the presence of nasal foramina, a posterodorsal process of the coronoid of the lower jaw, the present of caniniform successional teeth at the front of the jaws, the presence of flanges on the posterior parts of teeth at back of the lower jaw, and an expanded radial condyle on the humerus. Like modern tuatara, members of Sphenodontinae were likely generalists with a carnivorous/insectivorous diet.
Homoeosaurus is an extinct genus of rhynchocephalian reptile, known from the Late Jurassic-earliest Cretaceous of Europe, with specimens being reported from France, England and Germany. Several species have been described within the genus, based on varying proportions of the limb bones to the body length based on the presacral vertebrae. Specimen C.M.6438 of H. maximiliani from Germany has a total length of around 17 centimetres (6.7 in), with a skull length of about 1.7 centimetres (0.67 in). In comparison to other rhynchocephalians, the limbs are proportionally long. Recent studies have classified Homoeosaurus as a member of Neosphenodontia, with some studies including it as part of the clade Leptorhynchia, also including sapheosaurs, pleurosaurs, Kallimodon and Vadasaurus. Despite being found in aquatic deposits, it is suggested to have been terrestrial. It is thought to have been a carnivore/insectivore. One specimen was found as stomach contents of the fish Belonostomus.
Pleurosauridae is an extinct family of sphenodontian reptiles, known from the Jurassic of Europe. Members of the family had long-snake like bodies with reduced limbs that were adapted for aquatic life in marine environments. It contains two genera, Palaeopleurosaurus, which is known from the Early Jurassic (Toarcian) Posidonia Shale of Germany, as well as Pleurosaurus from the Late Jurassic of Germany and France. Paleopleurosaurus is more primitive than the later Pleurosaurus, with a skull similar to those of other sphenodontians, while that of Pleurosaurus is highly modified relative to other sphendontians. They likely swam via anguilliform locomotion. Vadasaurus and Derasmosaurus from the Late Jurassic and Early Cretaceous of Europe have been placed as part of this family in some studies, but lack the body elongation that typifies the other two genera.
Clevosaurus is an extinct genus of rhynchocephalian reptile from the Late Triassic and the Early Jurassic periods. Species of Clevosaurus were widespread across Pangaea, and have been found on all continents except Australia and Antarctica. Five species of Clevosaurus have been found in ancient fissure fill deposits in south-west England and Wales, alongside other sphenodontians, early mammals and dinosaurs. In regards to its Pangaean distribution, C. hadroprodon is the oldest record of a sphenodontian from Gondwana, though its affinity to Clevosaurus has been questioned.
Cynosphenodon is an extinct genus of rhynchocephalian in the family Sphenodontidae from the Middle Jurassic La Boca Formation of Tamaulipas, Mexico. It is known from a largely complete lower jaw and fragments of the upper jaw. It is suggested to be among the closest known relatives of the tuatara, with both being placed in the Sphenodontinae, which is supported by among other characters, the growth pattern of the teeth.
Opisthias is a genus of sphenodont reptile. The type species, Opisthias rarus, is known from the Late Jurassic (Kimmeridgian-Tithonian) of western North America.
Diphydontosaurus is an extinct genus of small rhynchocephalian reptile from the Late Triassic of Europe. It is the most primitive known member of Sphenodontia.
Sapheosaurus was an extinct genus of Late Jurassic sphenodont. Its skull was longer and narrower than that of Homoeosaurus. It was classified as a genus of sapheosaur by Michael Benton in 1985. It reached a length of 70 cm from snout to tail. Sapheosaurus belongs to the clade Sapheosauridae, that also includes other taxa like Kallimodon. It is believed to be one of two aquatic sphenodont lineages, with Pleurosauridae being the other.
Gephyrosauridae is an extinct family of rhynchocephalians that lived in the Late Triassic and Early Jurassic. They are generally considered to be rhynchocephalians that lie outside of Sphenodontia, but in some analyses they are recovered as more closely related to squamates than to sphenodontians.
Gephyrosaurus is a genus of early rhynchocephalian first described and named in 1980 by Susan E. Evans. They are distantly related to the extant Sphenodon with which they shared a number of skeletal features including a large tooth row along the side of the palatine bone and posterior process of the dentary bone. The type species, G. bridensis, lived during Early Jurassic in Wales, UK. Whiteside & Duffin (2017) described the second species, G. evansae, known from a partial maxilla recovered from Late Triassic (Rhaetian) fissure fills in Carboniferous Limestone in Somerset. Gephyrosaurus, other potential gephyrosaurids and Wirtembergia are the only rhynchocephalians to lie outside Sphenodontia in modern definitions of the group, and have been found to be more closely related to squamates in some phylogenetic analyses.
Oenosaurus is an extinct genus of sphenodontian reptile from the Late Jurassic (Tithonian) aged Mörnsheim Formation of Germany.
Sphenotitan is an extinct genus of rhynchocephalian reptile, known from the Late Triassic (Norian) Quebrada del Barro Formation of Argentina. It is the earliest known member of the herbivorous Elienodontinae, and the only one known from the Triassic. It was a large-sized sphenodontian, with an estimated skull length of over 10 centimetres (3.9 in). The skull is roughly triangular in shape, and had large upper temporal fenestrae. The region of the skull in front of the eye socket is short. The premaxillae forms beak, with a cutting edge similar to a chisel. The teeth of Sphenotitan, like other elienodontines, were large and wide, and designed for shredding vegetation, with blade-like palatal teeth on the roof of the mouth.
Sphenovipera jimmysjoyi is an extinct species of sphenodontian dated from the Middle Jurassic. If was discovered in the lower part of the La Boca Formation located in Tamaulipas, Mexico. Only the lower jaw of this organism has been discovered and studied. It is possibly the only species of rhynchocephalian yet discovered to show evidence of venom delivery.
Priosphenodon is an extinct, large herbiviorous eilenodontine rhynchocephalian known from the mid-Cretaceous (Albian-Turonian) of Argentina. It is one of the largest known sphenodontians.
Opisthodontia is a proposed clade of sphenodontian reptiles, uniting Opisthias from the Late Jurassic-earliest Cretaceous of Europe and North America with the Eilenodontinae, a group of herbivorous sphenodontians known from the Late Triassic to Late Cretaceous.
Vadasaurus is an extinct genus of rhynchocephalian closely related to the aquatic pleurosaurids. Although this genus was not as specialized as the eel-like pleurosaurs for aquatic life, various skeletal features support the idea that it had a semiaquatic lifestyle. The type species, Vadasaurus herzogi, was described and named in 2017. It was discovered in the Solnhofen Limestone in Germany, which is dated to the Late Jurassic. The generic name "Vadasaurus" is derived from "vadare", which is Latin for "to go" or "to walk forth", and "saurus", which means "lizard". "Vadare" is the root of the English word "wade", which is the reason it was chosen for this genus, in reference to its perceived semiaquatic habits. The specific name, "herzogi", refers to Werner Herzog, a Bavarian filmmaker.
Clevosaurs are an extinct group of rhynchocephalian reptiles from the Triassic and Jurassic periods.
Eilenodontinae are an extinct clade of reptiles belonging to Sphenodontia. They are either considered a subgroup of Opisthodontia, or Sphenodontidae. They had deep jaws with broad, closely packed teeth with thick enamel and noticeable wear facets. They were likely herbivorous, and probably chewed with a proal movement, with food shredded between the edges of opposing wear facets. Members of the group are known from South America, North America and Europe. The earliest known member of the group, Sphenotitan, is known from the Late Triassic of South America. while the youngest members are known from the early Late Cretaceous of South America. The group contains some of the largest known sphenodontians, with Priosphenodon suggested to be the largest known non aquatic sphenodontian, with an estimated body length of over 1 metre (3.3 ft).
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