Rhynchocephalia

Tuatara were originally classified as agamid lizards when they were first described by John Edward Gray in 1831. They remained misclassified until 1867, when Albert Günther of the British Museum noted features similar to birds, turtles, and crocodiles. He proposed the order Rhynchocephalia (from Ancient Greek ('''') 'beak' and ('''') 'head', meaning "beak head") for the tuatara and its fossil relatives. Sphenodon is derived from Ancient Greek ('''') 'wedge' and ('''') 'tooth'. Many disparately related species were subsequently added to the Rhynchocephalia, resulting in what taxonomists call a "wastebasket taxon". These include the superficially similar (both in shape and name) but unrelated rhynchosaurs, which lived in the Triassic. == Anatomy ==

Rhynchocephalia and their sister group Squamata (which includes lizards, snakes and amphisbaenians) belong to the superorder Lepidosauria, the only surviving taxon within Lepidosauromorpha. Squamates and rhynchocephalians have a number of shared traits (synapomorphies), including fracture planes within the tail vertebrae allowing caudal autotomy (loss of the tail when threatened), transverse cloacal slits, an opening in the pelvis known as the thyroid fenestra, the presence of extra ossification centres in the limb bone epiphyses, a knee joint where a lateral recess on the femur allows the articulation of the fibula, the development of a sexual segment of the kidney, and a number of traits of the feet bones, including a fused astralago-calcaneun and enlarged fourth distal tarsal, which creates a new joint, along with a hooked fifth metatarsal. Like some lizards, the tuatara possesses a parietal eye (also called a pineal eye or a third eye) covered by scales at the top of the head formed by the parapineal organ, with an accompanying hole in the skull roof enclosed by the parietal bones, dubbed the "pineal foramen", which is also present in fossil rhynchocephalians. The parietal eye detects light (though it is probably not capable of detecting movement or forming images), monitoring the day-night and seasonal cycles, helping to regulate the circadian rhythm, among other functions. While parietal eyes were widespread among early vertebrates, including early reptiles, they have been lost among most living groups. Unlike squamates, but similar to the majority of birds, the tuatara lacks a penis. This is a secondary loss, as a penis or squamate-like hemipenes were probably present in the last common ancestor of rhynchocephalians and squamates. '', which has an open lower temporal fenestra The complete lower temporal bar (caused by the fusion of the jugal and quadtrate/quadratojugal bones of the skull) of the tuatara, often historically asserted to be a primitive feature retained from earlier reptiles, is actually a derived feature among sphenodontians, with primitive lepidosauromorphs and many rhynchocephalians including the most primitive ones having an open lower temporal fenestra without a temporal bar. While often lacking a complete temporal bar, the vast majority of rhynchocephalians have a posteriorly directed process (extension) of the jugal bone. All known rhynchocephalians lack the splenial bone present in the lower jaw of more primitive reptiles, The dentition of most rhynchocephalians, including the tuatara, is described as acrodont, which is associated with the condition of the teeth being attached to the crest of the jaw bone, lacking tooth replacement and having extensive bone growth fusing the teeth to the jaws resulting in the boundary between the teeth and bone being difficult to discern. This differs from the condition found in most lizards (except acrodontans), which have pleurodont teeth which are attached to the shelf on the inward-facing side of the jaw, and are replaced throughout life. The teeth of the tuatara have no roots, though the teeth of some other rhynchocephalians possess roots (in addition, the precise technical meaning of term "acrodont" is somewhat ambiguous and the term is used with inconsistent meanings by different researchers). The most primitive rhynchocephalians have either pleurodont teeth or a combination of both pleurodont front and acrodont posterior teeth. In many derived sphenodontians, the premaxillary teeth at the front of the upper jaw are merged into a large chisel-like structure. and in combination with propalinal movement (back and forward motion of the lower jaw) allows for a shearing bite. Priosphenodon avelasi, one of the largest known sphenodontians The body size of rhynchocephalians is highly variable. The tuatara has an average total length of for females and males respectively. Clevosaurus sectumsemper has an estimated total length of , while large individuals of the largest known terrestrial sphenodontian, Priosphenodon avelasi reached total lengths of just over . The aquatic pleurosaurs reached lengths of up to . Most derived rhynchocephalians have the number of presacral vertebrae (the number of vertebrae forward of the sacrum) typically around 23-25, though the number is much greater in pleurosaurs, where it reaches up to 57 in some individuals. Seven cervical vertebrae in the neck may be typical for rhynchocephalians as it is for the tuatara. The tuatara retains a proatlas bone between the atlas (the first neck vertebra) and the skull, which has been lost in squamates. Its presence in fossil rhynchocephalians is unclear. Rhynchocephalians typically have amphicoelous vertebral centra (both faces are concave) on their presacral vertebrae. The tuatara has among the highest known ages of sexual maturity among reptiles, at around 9 to 13 years of age, and has a high longevity in comparison to lizards of similar size, Such a late onset of sexual maturity and longevity may have or not have been typical of extinct rhynchocephalians. == Classification ==

'' from the Late Jurassic of Germany While the grouping of Rhynchocephalia is well supported, the relationships of many taxa to each other are uncertain, varying substantially between studies. In modern cladistics, the clade Sphenodontia includes all rhynchocephalians other than Wirtembergia, as well as Gephyrosaurus and other gephyrosaurids. Gephyrosaurids have been found as more closely related to squamates in some analyses. In 2018, two major clades within Sphenodontia were defined, the infraorder Eusphenodontia which is defined by the least inclusive clade containing Polysphenodon, Clevosaurus hudsoni and Sphenodon, which is supported by the presence of three synapomorphies, including the presence of clearly visible wear facets on the teeth of the dentary or maxilla, the premaxillary teeth are merged into a chisel like structure, and the palatine teeth are reduced to a single tooth row, with the presence of an additional isolated tooth. The unranked clade Neosphenodontia is defined as the most inclusive clade containing Sphenodon but not Clevosaurus hudsoni, which is supported by the presence of six synapomorphies, including the increased relative length of the antorbital region of the skull (the part of the skull forward of the eye socket), reaching 1/4 to 1/3 of the total skull length, the posterior (hind) edge of the parietal bone is only slightly curved inward, the parietal foramen is found at the same level or forward of the anterior border of the supratemporal fenestra (an opening of the skull), the palatine teeth are further reduced from the condition in eusphenodontians to a single lateral tooth row, the number of pterygoid tooth rows are reduced to one or none, and the posterior border of the ischium is characterised by a distinctive process. In 2021 the clade Acrosphenodontia was defined, which is less inclusive than Sphenodontia and more inclusive than Eusphenodontia, and includes all sphenodontians with fully acrodont dentition, excluding basal partially acrodont sphenodontians. In 2022 the extinct clade Leptorhynchia was defined, including a variety of neosphenodontians, at least some of which were aquatically adapted, characterised by the elongation of the fourth metacarpal, the presence of a posterior process on the ischium, and the antorbital region of the skulls is between a third and a quarter of the total skull length. The clade Opisthodontia has been used for the grouping of all sphenodontians more closely related to Priosphenodon (a member of Eilenodontinae) than to Sphenodon. Not all studies use this clade, as some studies have found the scope of the clade to be identical to Eilenodontinae. The family Sphenodontidae has been used to include the tuatara and its closest relatives within Rhynchocephalia. However the grouping has lacked a formal definition, with the included taxa varying substantially between analyses. The closest relatives of the tuatara are placed in the clade Sphenodontinae, which are characterised by a completely closed temporal bar. The following is a cladogram of Rhynchocephalia after DeMar et al. 2022 (based on maximum parsimony, note that cladogram collapses into a polytomy under Bayesian analysis): }}}}}}|label1=Sphenodontia}} Clades and genera • †Wirtembergia • †Gephyrosauridae? • †Deltadectes • †Gephyrosaurus • †Penegephyrosaurus • †Bharatagama? • Sphenodontia • †Agriodontosaurus • †Diphydontosaurus • †Micromenodon • †Paleollanosaurus? • †Parvosaurus • †Pelecymala • †Whitakersaurus • Acrosphenodontia • †Godavarisaurus • †Planocephalosaurus • †Rebbanasaurus • †Theretairus • †Sphenocondor • Eusphenodontia • †Brachyrhinodon • †Colobops? • †Lanceirosphenodon • †Opisthiamimus • †Polysphenodon • †Clevosauridae • †Brachyrhinodon? • †Clevosaurus • †Polysphenodon? • †Microsphenodon • †Sigmala • †Trullidens • Neosphenodontia • †Derasmosaurus • †Lamarquesaurus • †Pamizinsaurus • †Tingitana • †Opisthodontia • †Alamitosphenos • †Eilenodontinae • Sphenodontidae • †Eilenodontinae? • †Eilenodon • †Kaikaifilusaurus • †Patagosphenos • †Priosphenodon • †Sphenotitan • †Toxolophosaurus • †Opisthias? • †Fraserosphenodon? • Sphenodontinae • Cynosphenodon • Tika • Sphenofontis • Navajosphenodon • Kawasphenodon • Notosphenos • Sphenodon • †Leptorhynchia • †Homoeosaurus • †Kallimodon • †Leptosaurus • †Vadasaurus • †Ankylosphenodon • †Sphenodraco • †Sapheosauridae • Oenosaurus • Piocormus • Sapheosaurus • †Pleurosauridae • Palaeopleurosaurus • Pleurosaurus • ?Derasmosaurus • ?Vadasaurus • ?Ankylosphenodon == Ecology ==

, an aquatically adapted sphenodontian from the Late Jurassic of GermanyThe fossil record of rhynchocephalians demonstrates that they were a diverse group that exploited a wide array of ecological niches. Like modern tuatara, extinct members of Sphenodontinae were likely generalists with a carnivorous/insectivorous diet. Amongst the most distinct rhynchocephalians are the pleurosaurs, known from the Jurassic of Europe, which were adapted for marine life, with elongated snake-like bodies with reduced limbs, with the specialised Late Jurassic genus Pleurosaurus having an elongated triangular skull highly modified from those of other rhynchocephalians. Pleurosaurs are thought to have been piscivores (consuming fish). with fish found as gut contents in one Kallimodon specimen. The sapheosaurids, such as Oenosaurus and Sapheosaurus from the Late Jurassic of Europe possess broad tooth plates unique amongst tetrapods, and are thought to have been durophagous, with the tooth plates being used to crush hard shelled organisms. though this interpretation has been questioned by other authors. The limb bone proportions and shape of the hand and foot bones of Sphenodraco from the Late Jurassic of Germany indicate that it was a primarily arboreal tree climbing animal, unlike the largely terrestrial tuatara. Other extinct rhynchocephalians with relatively long limbs such as Navajosphenodon and Homoeosaurus'' may also have exhibited climbing capabilities. == Evolutionary history ==

(left) and Clevosaurus cambrica'' (right) The timing of when Rhynchocephalia is estimated to have diverged from Squamata is disputed. Older estimates place the divergence between the Middle Permian and earliest Triassic, around 270 to 252 million years ago, The next earliest rhynchocephalian is Wirtembergia which is known from the Erfurt Formation near Vellberg in Southern Germany, dating to the Ladinian stage of the Middle Triassic, around 238-240 million years old. The earliest rhynchocephalians were small animals, but by the Late Triassic the group had evolved a wide range of body sizes. During the Jurassic, rhynchocephalians were the dominant group of small reptiles globally, reaching their apex of morphological diversity during this period, including specialised herbivorous and aquatic forms. The only record of Rhynchocephalians from Asia (excluding the Indian subcontinent, which was not part of Asia during the Mesozoic) are indeterminate remains of Clevosaurus from the Early Jurassic (Sinemurian) aged Lufeng Formation of Yunnan, China. Rhynchocephalians are noticeably absent from younger localities in the region, despite the presence of favourable preservation conditions. Rhynchocephalians remained diverse into the Late Jurassic, and were absent from North Africa by the early Late Cretaceous. The cause of the decline of Rhynchocephalia remains unclear, but has often been suggested to be due to competition with advanced lizards and mammals. Late Cretaceous South American sphenodontians are represented by both Eilenodontinae and Sphenodontidae (including Sphenodontinae). An indeterminate rhynchocephalian is known from a partial lower jaw of a hatchling from the latest Cretaceous or possibly earliest Paleocene Intertrappean Beds, in what was then the isolated landmass of Insular India, which appears to be an acrosphenodontian, possibly related to Godavarisaurus from the Jurassic of India. The youngest undoubted remains of rhynchocephalians outside of New Zealand are those of the sphenodontid Kawasphenodon peligrensis from the early Paleocene (Danian) of Patagonia approximately 64-63 million years ago, shortly after the Cretaceous–Paleogene extinction event. Indeterminate sphenodontine jaw fragments bearing teeth are known from the early Miocene (19–16 million years ago) St Bathans fauna, New Zealand, that are indistinguishable from those of the living tuatara. It is unlikely that the ancestors of the tuatara arrived in New Zealand via oceanic dispersal, and it is thought that they were already present in New Zealand when it separated from Antarctica between 80 and 66 million years ago. == References ==