Squamata

Squamata is the largest order of reptiles, comprising lizards, snakes and amphisbaenians (worm lizards), which are collectively known as squamates or scaled reptiles. With over 10,000 species, it is also the second-largest order of extant (living) vertebrates, after the perciform fish, and roughly equal in number to the Saurischia (one of the two major groups of dinosaurs). Members of the order are distinguished by their skins, which bear horny scales or shields. They also possess movable quadrate bones, making it possible to move the upper jaw relative to the neurocranium. This is particularly visible in snakes, which are able to open their mouths very wide to accommodate comparatively large prey. Squamata is the most variably sized order of reptiles, ranging from the 16 mm (0.63 in) dwarf gecko (Sphaerodactylus ariasae) to the 5.21 m (17.1 ft) green anaconda (Eunectes murinus) and the now-extinct mosasaurs, which reached lengths of over 14 m (46 ft). Among other reptiles, squamates are most closely related to the tuatara, which superficially resembles lizards. Squamates are a monophyletic sister group to the rhynchocephalians, members of the order Rhynchocephalia. The only surviving member of Rhynchocephalia is the tuatara. Squamata and Rhynchocephalia form the subclass Lepidosauria, which is the sister group to Archosauria, the clade that contains crocodiles and birds, and their extinct relatives. Fossils of rhynchocephalians first appear in the Early Triassic, meaning that the lineage leading to squamates must have also existed at the time. Scientists believe crown group squamates probably originated in the Early Jurassic based on the fossil record. The first fossils of geckos, skinks and snakes appear in the Middle Jurassic. Other groups like iguanians and varanoids appeared in the Cretaceous. Polyglyphanodontians, a distinct clade of lizards, and mosasaurs, a group of predatory marine lizards that grew to enormous sizes, also appeared in the Cretaceous. Squamates suffered a mass extinction at the Cretaceous–Paleogene (K–PG) boundary, which wiped out polyglyphanodontians, mosasaurs and many other distinct lineages. The relationships of squamates is debatable. Although many of the groups originally recognized on the basis of morphology are still accepted, our understanding of their relationships to each other has changed radically as a result of studying their genomes. Iguanians were long thought to be the earliest crown group squamates based on morphological data, however, genetic data suggests that geckoes are the earliest crown group squamates. Iguanians are now united with snakes and anguimorphs in a clade called Toxicofera. Genetic data also suggests that the various limbless groups; snakes, amphisbaenians and dibamids, are unrelated, and instead arose independently from lizards. A study in 2018 found that Megachirella, an extinct genus of lepidosaur that lived about 240 million years ago during the Middle Triassic, was a stem-squamate, making it the oldest known squamate. The phylogenetic analysis was conducted by performing high-resolution microfocus X-ray computed tomography (micro-CT) scans on the fossil specimen of Megachirella to gather detailed data about its anatomy. This data was then compared with a phylogenetic dataset combining the morphological and molecular data of 129 extant and extinct reptilian taxa. The comparison revealed Megachirella had certain features that are unique to squamates. The study also found that geckos are the earliest crown group squamates not iguanians. The male members of the group Squamata have hemipenes, which are usually held inverted within their bodies, and are everted for reproduction via erectile tissue like that in the human penis. Only one is used at a time, and some evidence indicates that males alternate use between copulations. The hemipenis has a variety of shapes, depending on the species. Often it bears spines or hooks, to anchor the male within the female. Some species even have forked hemipenes (each hemipenis has two tips). Due to being everted and inverted, hemipenes do not have a completely enclosed channel for the conduction of sperm, but rather a seminal groove that seals as the erectile tissue expands. This is also the only reptile group in which both viviparous and ovoviviparous species are found, as well as the usual oviparous reptiles. Some species, such as the Komodo dragon, can reproduce asexually through parthenogenesis. There have been studies on how sexual selection manifests itself in snakes and lizards. Snakes use a variety of tactics in acquiring mates. Ritual combat between males for the females they want to mate with includes topping, a behavior exhibited by most viperids, in which one male will twist around the vertically elevated fore body of its opponent and forcing it downward. It is common for neck biting to occur while the snakes are entwined. Parthenogenesis is a natural form of reproduction in which the growth and development of embryos occur without fertilization. Agkistrodon contortrix (copperhead snake) and Agkistrodon piscivorus (cotton mouth snake) can reproduce by facultative parthenogenesis. That is, they are capable of switching from a sexual mode of reproduction to an asexual mode. The type of parthenogenesis that likely occurs is automixis with terminal fusion (see figure), a process in which two terminal products from the same meiosis fuse to form a diploid zygote. This process leads to genome wide homozygosity, expression of deleterious recessive alleles and often to developmental abnormalities. Both captive-born and wild-born A. contortrix and A. piscivorus appear to be capable of this form of parthenogenesis.