Rhynchocephalia

Rhynchocephalia (/ˌrɪŋkoʊsɪˈfeɪliə/; lit. 'beak-heads') is an order of lizard-like reptiles that includes only one living species, the tuatara (Sphenodon punctatus) 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 244 million years ago, and they had achieved global distribution by the Early Jurassic. Most rhynchocephalians belong to the suborder 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.

Rhynchocephalians are distinguished from squamates by a number of traits, including the retention of rib-like gastralia bones in the belly, as well as most rhynchocephalians having acrodont teeth that are fused to the crests of the jaws (the latter also found among a small number of modern lizard groups like agamids).

Once representing the world's dominant group of small reptiles, many of the niches occupied by lizards today were held by rhynchocephalians during the Triassic and Jurassic. Rhynchocephalians underwent a great decline during the Cretaceous, and they had disappeared almost entirely by the beginning of the Cenozoic. While the modern tuatara is primarily insectivorous and carnivorous, the diversity of the group also included the herbivorous eilenodontines, and there were other rhynchocephalians with specialised ecologies like the durophagous sapheosaurs. There were even successful groups of aquatic sphenodontians, such as the elongate-bodied pleurosaurs.

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 ῥύγχος (rhúnkhos) 'beak' and κεφαλή (kephalḗ) 'head', meaning "beak head") for the tuatara and its fossil relatives. In 1925, Samuel Wendell Williston proposed the Sphenodontia to include only tuatara and their closest fossil relatives. Sphenodon is derived from Ancient Greek σφήν (sphḗn) 'wedge' and ὀδούς (odoús) '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. Studies in the 1970s and 1980s demonstrated that many rhynchosaurs were unrelated, with computer-based cladistic analysis conducted in the 1980s providing a robust diagnosis for the definition of the group.

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.

Rhynchocephalians are distinguished from squamates by a number of traits, including the retention of gastralia (rib-like bones present in the belly of the body, ancestrally present in tetrapods and also present in living crocodilians). 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.