| Clevosaurs Temporal range: | |
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| Artist's illustration of Clevosaurus hadroprodon | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Order: | Rhynchocephalia |
| Suborder: | Sphenodontia |
| Family: | † Clevosauridae Bonaparte & Sues, 2006 |
| Genera | |
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Clevosaurs are an extinct group of rhynchocephalian reptiles from the Triassic and Jurassic periods. [1]
Although members of this group have been known since 1910, only recently has the group received a formal name. In the late 1990s, Victor-Hugo Reynoso established that three particular genera of Sphenodontia ( Clevosaurus, Brachyrhinodon , and Polysphenodon ) were closely related to each other. He gave the informal name "clevosaurs" to these three genera, after the most numerous and well-known genus, Clevosaurus. He considered clevosaurs to be members of the family Sphenodontidae, the family of rhynchocephalians containing the only living member of the order, the tuatara (Sphenodon).
In 2006, Bonaparte and Sues finally gave "clevosaurs" a formal name and taxonomic rank as the family Clevosauridae. They defined Clevosauridae as the last common ancestor of Clevosaurus, Brachyrhinodon, and Polysphenodon, and all of its descendants. [1] In 2015, the definition of this family was revised to be "all taxa more closely related to Clevosaurus than to Sphenodon". [2] However, the erection of this family conflicts with their position within Sphenodontidae, as a taxonomic family cannot be within another family. Sources which use Sphenodontidae as a wide group of sphenodontians do not use the term Clevosauridae, instead continuing to use the informal term 'clevosaurs'. On the other hand, sources which use Clevosauridae do not use Sphenodontidae. Some do not use either family, instead opting for 'clevosaurs' and 'advanced sphenodontians'. [3]
Clevosaurs were among the first major groups of sphenodontians to evolve, and had a worldwide distribution in the Late Triassic and early Jurassic. Clevosaurus was particularly widespread and diverse, surviving the Triassic-Jurassic extinction and being known from numerous species.
Adult clevosaurs are notable among sphenodontians for their short, boxy snouts. The antorbital region of the skull (the portion in front of the eyes) only occupies a quarter of the length of the entire skull in most clevosaurs, although a few species of Clevosaurus reacquire a slightly longer skull. Like other rhynchocephalians, they possessed two pairs of large holes called temporal fenestrae in the back part of the skull. The lower temporal fenestrae (on the sides of the skull) are very large in most clevosaurs, about a quarter the length of the skull. All clevosaurs have very long jugal bones which extend back as far as the squamosal bones in the back of the head, forming the entire upper edge of their lower temporal fenestrae in the process. [2]
Like other sphenodontians, clevosaurs had several rows of teeth on the roof of the mouth. Their teeth were acrodont, meaning that they grew directly from the bone rather than from tooth sockets. The large outermost row of teeth were attached to the maxillae bones while the small innermost teeth were clustered in rows on the pterygoid bones. Between the maxillary and pterygoid teeth, clevosaurs characteristically had one row of large teeth on each palatine bone, as well as an additional isolated tooth at the inner front corner of each palatine. [2]
Clevosaurs also had a row of teeth on the edge of their dentaries (lower jaws). In young individuals, these teeth were spike-like, well-adapted for consuming insects and other invertebrates. However, as individual clevosaurs grew older, their jaws became shorter and more robust. In addition, both their maxillary and dentary teeth wear down into a sharp cutting edge, creating a "beak"-like jaw structure somewhat similar to the jaws of modern Uromastyx lizards. It is likely that adult clevosaurs may have been omnivorous or herbivorous, similar to Uromastyx in ecology. [4]
Below is a cladogram of the relationships within Clevosauridae based on the phylogenetic analysis of Hsiou et al. (2015): [2]
| Clevosauridae | |||||||||||||
"Clevosaurus" latidens was recovered outside of Clevosauridae, as the sister taxon of Opisthodontia. [2]
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 also sometimes considered part of this family as the sister group to Sphenodontinae. Sphenodontines first appeared during the Early Jurassic, and are characterised by a complete lower temporal bar caused by the fusion of the quadrate/quadratojugal and the jugal, which was an adaptation for reducing stress in the skull during hard biting. Like modern tuatara, members of Sphenodontinae were likely generalists with a carnivorous/insectivorous diet.
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.
Eilenodon is an extinct genus of rhynchocephalian reptile from the Late Jurassic Morrison Formation of western North America, present in stratigraphic zone 4. The only known species of this genus was Eilenodon robustus. It was a member of a group of rhynchocephalians called the eilenodontines, which were large, herbivorous members of Rhynchocephalia, the order of reptiles which contains the modern tuatara (Sphenodon). The generic name "Eilenodon" is Greek for "packed teeth", in reference to its closely packed teeth. The specific name, "robustus", refers to the strong build of the jaws.
Diphydontosaurus is an extinct genus of small rhynchocephalian reptile from the Late Triassic of Europe. It is the most primitive known member of Sphenodontia.
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.
Zapatadon is an extinct genus of sphenodontid reptile from the end of the Early Jurassic in the lower part of La Boca Formation of Tamaulipas, Mexico. Is known from a nearly complete skull with mandible of a post-hatchling individual, and is one of the smallest skulls between the sphenodontians, with an estimated total length of 11.3 millimetres, a bit smaller than the hatchling individuals observed in the modern tuatara (Sphenodon); features like the oblique mandibular symphysis suggests that the holotype is from an individual in a relatively mature stage of ontogenic development. Zapatadon is diagnosed by their hatchling tooth series located in a depression in the anterior part of the dentary bone, the prefrontal bone surrounding the dorsal process of the maxilla and the broad jugal that extends over the maxillary suborbital process, been almost excluded of the orbit.
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.
Whitakersaurus is a genus of sphenodontid rhynchocephalian reptile dated to be late Triassic in age and is from the Ghost Ranch fossil quarry in New Mexico, USA. It is named after the discoverer of the Ghost Ranch quarry, George O. Whitaker. The fossil was described in 2007.
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.
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 and thus cannot be assigned its own family. They were fairly advanced rhynchocephalians which may have had semiaquatic habits.
Colobops is a genus of reptile from the Late Triassic of Connecticut. Only known from a tiny skull, this reptile has been interpreted to possess skull attachments for very strong jaw muscles. This may have given it a very strong bite, despite its small size. However, under some interpretations of the CT scan data, Colobops's bite force may not have been unusual compared to other reptiles. The generic name, Colobops, is a combination of κολοβός, meaning shortened, and ὤψ, meaning face. This translation, "shortened face", refers to its short and triangular skull. Colobops is known from a single species, Colobops noviportensis. The specific name, noviportensis, is a latinization of New Haven, the name of both the geological setting of its discovery as well as a nearby large city. The phylogenetic relations of Colobops are controversial. Its skull shares many features with those of the group Rhynchosauria, herbivorous archosauromorphs distantly related to crocodilians and dinosaurs. However, many of these features also resemble the skulls of the group Rhynchocephalia, an ancient order of reptiles including the modern tuatara, Sphenodon. Although rhynchosaurs and rhynchocephalians are not closely related and have many differences in the skeleton as a whole, their skulls are remarkably similar. As Colobops is only known from a skull, it is not certain which one of these groups it belonged to. Pritchard et al. (2018) interpreted it as a basal rhynchosaur, while Scheyer et al. (2020) reinterpreted it as a rhynchocephalian.
Fraxinisaura is an extinct genus of basal lepidosauromorph reptile known from the Middle Triassic of Germany. The only known species is Fraxinisaura rozynekae. It possessed an elongated snout, unique features of the teeth, and an ilium which was intermediate in orientation between sphenodontians and squamates. Based on characteristics of the maxilla, it is considered a close relative of Marmoretta from the Middle Jurassic of the United Kingdom, resolving a ghost lineage between that genus and other Triassic basal lepidosauromorphs.
Eilenodontinae are an extinct clade of reptiles belonging to Sphenodontia, related to the modern tuataras. 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.
Kawasphenodon is an extinct genus of sphenodontian reptile, known from the Late Cretaceous and Paleocene of Patagonia in South America. The type species, K. expectatus, was described in 2005 from jaw fragments found in late Campanian aged sediments in the Los Alamitos Formation, the jaw when complete was estimated to be 11 cm long, making it among the largest known sphenodontians. A second species, K. peligrensis, around 1/3 the size of the type species, was described in 2014 also from jaw fragments in early Paleocene (Danian) sediments of the Salamanca Formation, making it the youngest known definitive representative of Rhynchocephalia outside of New Zealand. In the original description, it was found to be a member of Sphenodontidae, in some other subsequent analyses it was found to be a member of Opisthodontia. A 2020 analysis of rhyncocephalian relationships found it to be outside Opisthodontia, and instead a member of the Sphenodontinae as the closest known relative of the tuatara, with an estimated divergence between the two genera in the Early Cretaceous. Other subsequent studies have endorsed its placement as a member of Sphenodontidae. Like most other rhynchocephalians, the teeth are acrodont, with a deep dentary, and it probably had an omnivorous habit.
Taytalura is an extinct genus of lepidosauromorph reptile from the Late Triassic of Argentina. It contains a single species, Taytalura alcoberi, which is based on a well-preserved skull from the fossiliferous Ischigualasto Formation. As a lepidosauromorph, Taytalura is a distant relative of modern lepidosaurs such as sphenodontians and squamates. Taytalura did not belong to any group of modern lepidosaurs, since it bears unique features, such as unfused bones in the skull roof and teeth which all sit loosely in a deep groove without sockets. Regardless, Micro-CT scanning reveals features of the skull previously only seen in rhynchocephalians. This suggests that the ancestral condition of the skull in lepidosaurs was more similar to sphenodonts than to squamates.
Microsphenodon is an extinct genus of sphenodontian from the Late Triassic of Brazil. The type species is Microsphenodon bonapartei. It is a small sphenodontian with a skull roughly 20 mm long, and represents a unique mosaic of characteristics shared by both early diverging sphenodontians and eusphenodontian characters. Specimens of this taxon were first identified by Bonaparte and Sues (2006) but were misidentified as juvenile Clevosaurus brasiliensis, characteristics such as differences in the configuration of the teeth on the palate, and the unique form of tooth implantation seen in C. brasiliensis, helped differentiate these two co-occurring sphenodontian taxa.
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