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Squamata

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Squamata

Squamata (/skwæˈmtə/, Latin squamatus, 'scaly, having scales') is the largest order of reptiles; most members of which are commonly known as lizards, with the group also including snakes. With over 11,991 species, it is also the second-largest order of extant (living) vertebrates, after the perciform fish. Squamates are distinguished by their skins, which bear horny scales or shields, and must periodically engage in molting. They also possess movable quadrate bones, making possible movement of the upper jaw relative to the neurocranium. This is particularly visible in snakes, which are able to open their mouths very widely to accommodate comparatively large prey. Squamates are the most variably sized living reptiles, ranging from the 16 mm (0.63 in) dwarf gecko (Sphaerodactylus ariasae) to the 6.5 m (21 ft) reticulated python (Malayopython reticulatus). The now-extinct mosasaurs reached lengths over 14 m (46 ft).

Among other reptiles, squamates are most closely related to the tuatara, the last surviving member of the once diverse Rhynchocephalia, with both groups being placed in the superorder Lepidosauria.

Squamates are a monophyletic sister group to the rhynchocephalians, members of the order Rhynchocephalia. The only surviving member of the Rhynchocephalia is the tuatara. Squamata and Rhynchocephalia form the superorder Lepidosauria, which is the sister group to the 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.

A study in 2018 found that Megachirella, an extinct genus of lepidosaurs 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. These data were 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. However, a 2021 study found the genus to be a lepidosaur of uncertain position, in a polytomy with Squamata and Rhynchocephalia.

In 2022, the extinct genus Cryptovaranoides was described from the Late Triassic (Rhaetian age) of England as a highly derived squamate belonging to the group Anguimorpha, which contains many extant lineages such as monitor lizards, beaded lizards and anguids. The presence of an essentially modern crown group squamate so far back in time was unexpected, as their diversification was previously thought to have occurred during the Jurassic and Cretaceous. A 2023 study found that Cryptovaranoides most likely represents an archosauromorph with no apparent squamate affinities, though the original describers maintained their original conclusion that this taxon represents a squamate. The oldest unambiguous fossils of Squamata date to the Bathonian age of the Middle Jurassic of the Northern Hemisphere, with the first appearance of many modern groups, including snakes, during this period.

Scientists believe crown group squamates probably originated in the Early Jurassic based on the fossil record, with the oldest unambiguous fossils of squamates dating to the Middle Jurassic. Squamate morphological and ecological diversity substantially increased over the course of the Cretaceous, including the appeance of groups like iguanians and varanoids, and true snakes. Polyglyphanodontia, an extinct 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 are debatable. Although many of the groups originally recognized on the basis of morphology are still accepted, 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, but genetic data suggest that geckos are the earliest crown group squamates. Iguanians are now united with snakes and anguimorphs in a clade called Toxicofera. Genetic data also suggest that the various limbless groups – snakes, amphisbaenians, and dibamids – are unrelated, and instead arose independently from lizards.

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 mammalian 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. The eggs in oviparous species have a parchment-like shell. The only exception is found in blind lizards and three families of geckos (Gekkonidae, Phyllodactylidae and Sphaerodactylidae), where many lay rigid and calcified eggs. Some species, such as the Komodo dragon, can reproduce asexually through parthenogenesis.

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