Opisthodontia (reptile)

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Opisthodonts
Temporal range: Late Triassic–Late Cretaceous
Priosphenodon NT small.jpg
Priosphenodon , a large eilenodontine opisthodont
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Rhynchocephalia
Suborder: Sphenodontia
Clade: Opisthodontia
Apesteguia & Novas, 2003
Genera

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.

Contents

Description

Teeth and diet

Like other sphenodonts, opisthodonts had acrodont teeth which grew directly from the bone. They had one row of teeth on the lower jaw and two rows on the roof of the mouth. When processing food, their mandibular teeth would have slid between the outer (maxillary) teeth and inner (palatine) teeth. [1] Some opisthodonts, such as Sphenotitan , also had clusters of small teeth on the pterygoid at the center of the mouth roof. Opisthodont teeth were wide, numerous, and tightly-packed for grinding and shredding tough plant matter. Although wide shredding teeth are also known in a few other sphenodontians, such as Clevosaurus and Pelecymala , the most diverse and long-lasting group of herbivorous rhynchocephalians were the opisthodonts. [2] Some more generalized opisthodonts, such as Opisthias , were probably more capable of omnivory than the advanced eilenodonts. [3]

The mandibular teeth of opisthodonts were characteristically edged with forward-pointing flanges. Although Sphenotitan only had flanges on the medial (inside) edge, other eilenodonts had flanges on the lateral (outside) edge as well, making their teeth anteriorly concave and overlapping. The maxillary teeth, on the other hand, had long flanges aligned in a posteromedial-anterolateral direction. [4]

Opisthodonts lacked dental regionalization, meaning that all of their teeth had the same form and they did not have caniform or hatchling teeth like other sphenodonts. [2] Despite having had thicker enamel to resist wearing down their teeth, opisthodonts have often been found with their teeth significantly worn away, especially towards the front of the jaw. [1] The tip of the lower jaw is completely toothless in opisthodonts. Their jaws were also deep (particularly in eilenodonts) to counter stresses which would have occurred during food processing. [1]

Lower jaws of Sphenodon and several opisthodonts. Opisthodont jaws.png
Lower jaws of Sphenodon and several opisthodonts.

Characteristic features

Opisthodonts can be characterized by the following synapomorphies: [2] [5]

In addition, most opisthodonts also had dental ridges (or crests) on their mandibular teeth. The most advanced opisthodonts belong to the subfamily (or tribe) Eilenodontinae (or Eilenodontini). Most eilenodonts were large, stockily built members of the clade, with massive jaws, low and overlapping teeth, and a large hooked "beak"(a rhynchocephalian feature). Their skulls were reminiscent of rodent skulls, leading some to presume that they were gregarious burrowers which fed on tough vegetation. [4] The largest known terrestrial rhynchocephalian was an eilenodont, Priosphenodon avelasi.

Classification

First defined as a family by Apesteguia and Novas in 2003, Opisthodontia has generally been considered a well-supported clade, especially compared to other groups within Rhynchocephalia, which often shift in evolutionary position between studies. In particular, they are generally considered to be fairly close relatives of the family Sphenodontidae, which includes Sphenodon. Both opisthodonts and sphenodontids share the ability to move their jaw in a forward-to-back chewing motion, earning them the informal name "eupropalinal sphenodonts" as a result. Other possibly herbivorous rhynchocephalians, such as Pelecymala and Ankylosphenodon, have sometimes been suggested to be very close relatives (or basal members) of Opisthodontia in some studies, although these hypotheses are not well supported. [2] Ankylosphenodon is considered to be placed within Sphenodontidae, while Pelecymala is inconsistent in placement but may be a quite basal rhynchocephalian. [6] "Clevosaurus" latidens, which had previously been referred to Pelecymala and later Clevosaurus , was suggested to be an opisthodont in 2013, a hypothesis which was confirmed through a 2015 phylogenetic analysis. This same analysis found that Opisthias and Kawasphenodon formed a clade at the base of Opisthodontia, with Sphenotitan being a basal eilenodont. [7] In 2018, "Clevosaurus" latidens was named as the new genus Fraserosphenodon , and placed as the most basal opisthodont. [6]

Distribution

Skulls of Priosphenodon individuals of various ages Ontogenia8.jpg
Skulls of Priosphenodon individuals of various ages

Opisthodonts were numerous at times, with Opisthias having a practically worldwide distribution in the Jurassic and Priosphenodon being known from multiple specimens from the early Cretaceous of Argentina.

The oldest opisthodonts include Sphenotitan and Fraserosphenodon, from the Norian and Rhaetian (late Triassic) of Argentina and England, respectively. Fraserosphenodon is considered to be the most basal opisthodont. [6] Although Sphenotitan was also once believed to be the most basal opisthodont, it also shows many similarities with eilenodonts, and may have been the most basal member of that subfamily. [7] The oldest known non-Sphenotitan eilenodonts lived during the Jurassic and probably became extinct near the end of the Cretaceous. Opisthodonts as a whole survived a major extinction of Laurasian rhynchocephalians in the early Cretaceous as well as the K/T extinction at the end of the Cretaceous which led to the downfall of non-avian dinosaurs. The last known species of opisthodont was Kawasphenodon peligrensis , a small patagonian member of the group which lived during the Paleocene. The last of the opisthodonts probably became extinct due to the cooling climate of the late Eocene. [3]

Related Research Articles

<span class="mw-page-title-main">Rhynchocephalia</span> Order of reptiles

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.

<span class="mw-page-title-main">Sphenodontidae</span> Family of reptiles

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.

<i>Clevosaurus</i> Extinct genus of reptiles

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.

<i>Opisthias</i> Extinct genus of reptiles

Opisthias is a genus of sphenodont reptile. The type species, Opisthias rarus, is known from the Late Jurassic (Kimmeridgian-Tithonian) Morrison Formation of western North America, present in stratigraphic zones 2 and 4–6.

<i>Eilenodon</i> Extinct genus of reptiles

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.

Toxolophosaurus was a sphenodont which lived in North America during the Early Cretaceous.

<i>Sphenotitan</i> Extinct genus of reptiles

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.

<i>Sphenovipera</i> Extinct genus of reptiles

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.

<i>Priosphenodon</i> Extinct genus of reptiles

Priosphenodon is an extinct rhynchocephalian known from the Early to Late Cretaceous of Argentina. It is believed to have been a fairly large herbivore, having a longer snout than modern tuatara, with teeth specialised for shearing plant matter, like other members of Eilenodontinae. At slightly over 1 metre (3.3 ft) in total length, Priosphenodon is the largest known terrestrial sphenodontian. The teeth are densely packed with a cone-in-cone structure, and have prismatic enamel structure similar to those of mammals and the lizard Uromastyx, which was likely an adaptation to wear resistance in the absence of tooth replacement. Two species are known, P. avelesi and P. minimus. The latter has also been considered a part of the separate genus Kaikaifilusaurus.

<span class="mw-page-title-main">Clevosaurs</span> Family of reptiles

Clevosaurs are an extinct group of rhynchocephalian reptiles from the Triassic and Jurassic periods.

<span class="mw-page-title-main">Sapheosaur</span> Extinct group of reptiles

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.

<i>Lamarquesaurus</i> Extinct genus of reptiles

Lamarquesaurus is an extinct genus of sphenodontian from the Late Cretaceous (Campanian) of Patagonia. It is known from a single species, Lamarquesaurus cabazai. This genus and species is represented by MML-PV-42, a well-preserved right maxillary bone. This bone was found at the Cerro Tortuga site near Lamarque, Argentina, which preserves fossils from the Cretaceous Allen Formation. The discoverer of the Cerro Tortuga locality, Tito Cabaza, is the namesake of the species.

Kaikaifilusaurus is an extinct genus of rhynchocephalians in the family Sphenodontidae from the Late Cretaceous of South America. Fossils of the genus were found in Cenomanian sediments of the Candeleros Formation and Turonian layers of the Huincul Formation, both of the Neuquén Basin and the Albian strata of the Cerro Barcino Formation in the Cañadón Asfalto Basin, all in Patagonia, Argentina. The genus contains two species, K. minimus and the type species K. calvoi.

<span class="mw-page-title-main">Eilenodontinae</span> Subfamily of reptiles (fossil)

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.

<i>Microsphenodon</i> LAW, Constitution, History, Policy

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.

Fraserosphenodon is an extinct genus of sphenodontian from the Late Triassic of the United Kingdom. It contains a single species, Fraserosphenodon latidens.

Pelecymala is an extinct genus of sphenodontian reptile that lived in southwest England during the Triassic period. It has been recovered in recent studies as a primitive member of the group.

References

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