Squamata

This article is about the Squamata order of reptiles. For the Roman scale armour, see Lorica squamata.

Scaled reptiles Temporal range: Early Jurassic - Present, 199–0 Ma PreЄ Є O S D C P T J K Pg N
Eastern blue-tongued lizard
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Superorder:	Lepidosauria
Order:	Squamata Oppel, 1811
Subgroups
Dibamidae Gekkota Lacertoidea Scincomorpha Toxicofera Anguimorpha Iguania Ophidia
black: Range of Squamata

The Squamata, or the scaled reptiles, are the largest recent order of reptiles, comprising all lizards and snakes. With over 9,000 species, it is also the second-largest order of vertebrates, after the perciform fish. 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. They are 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 14 m (46 ft).

Among the other reptiles, squamates are most closely related to tuataras, which superficially resemble lizards.

Evolution

Slavoia darevskii, a fossil squamate

Squamates are a monophyletic sister group to the tuatara. The squamates and tuatara together are a sister group to crocodiles and birds, the extant archosaurs. Fossils of the squamate sister group, the Rhynchocephalia, appear in the Early Triassic, meaning that the lineage leading to squamates must have existed as well. Modern squamates probably originated in the mid Jurassic, when fossil relatives of geckos and skinks and snakes appear; other groups, including iguanians and varanoids, first appear in the Cretaceous period. Also appearing in the Cretaceous are the polyglyphanodonts, a lizard group of uncertain affinities, and the mosasaurs, a group of predatory, marine lizards that grew to enormous sizes. At the end of the Cretaceous, squamates suffered a major extinction at the K-T boundary which wiped out polyglyphanodonts, mosasaurs, and a number of other groups.

The relationships of squamates have been debated. 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 DNA. From morphological data, the iguanians were long thought to be the most ancient branch of the tree; however, studies of the DNA suggest that the geckos represent the most ancient branch. Iguanians are now united with snakes and anguimorphs in a group called the Toxicofera. DNA also suggests that the various limbless groups- snakes, amphisbaenians, and dibamids- are unrelated, and instead arose independently from lizards.

Reproduction

See also: Sexual selection in scaled reptiles

Trachylepis maculilabris skinks mating

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 actually reproduce asexually through parthenogenesis.

The Japanese striped snake has been studied in sexual selection

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.

Evolution of venom

Main article: Evolution of snake venom

See also: Venom

Recent research suggests that the evolutionary origin of venom may exist deep in the squamate phylogeny, with 60% of squamates placed in this hypothetical group called Toxicofera. Venom has been known in the clades Caenophidia, Anguimorpha, and Iguania, and has been shown to have evolved a single time along these lineages before the three groups diverged, because all lineages share nine common toxins. The fossil record shows the divergence between anguimorphs, iguanians, and advanced snakes dates back roughly 200 Mya to the Late Triassic/Early Jurassic. But the only good fossil evidence is from the Jurassic.

Snake venom has been shown to have evolved via a process by which a gene encoding for a normal body protein, typically one involved in key regulatory processes or bioactivity, is duplicated, and the copy is selectively expressed in the venom gland. Previous literature hypothesized that venoms were modifications of salivary or pancreatic proteins, but different toxins have been found to have been recruited from numerous different protein bodies and are as diverse as their functions.

Natural selection has driven the origination and diversification of the toxins to counter the defenses of their prey. Once toxins have been recruited into the venom proteome, they form large, multigene families and evolve via the birth-and-death model of protein evolution, which leads to a diversification of toxins that allows the ambush predators the ability to attack a wide range of prey. The rapid evolution and diversification is thought to be the result of a predator/prey evolutionary arms race, where both are adapting to counter the other.

Humans and squamates

Bites and fatalities

See also: Snakebite

Map showing the global distribution of snakebite morbidity

An estimated 125,000 people a year die from venomous snake bites. In the US alone, more than 8,000 venomous snake bites are reported each year.

Lizard bites, unlike venomous snake bites, are not fatal. The Komodo dragon has been known to kill people due to its size, and recent studies show it may have a passive envenomation system. Recent studies also show that the close relatives of the Komodo, the monitor lizards, all have a similar envenomation system, but the toxicity of the bites is relatively low to humans.

Conservation

Though they survived the most drastic changes in Earth's history, many squamate species are endangered now due to habitat loss, hunting and poaching, illegal wildlife trading, alien species being introduced to their habitats (which puts native creatures at risk through competition, disease, and predation), and many other unnecessary reasons. Because of this, some are in fact extinct, with Africa having the most extinct species of squamates. However, breeding programs and wildlife parks are trying to save many endangered reptiles from extinction. Many zoos, private hobbyists and breeders educate people about the importance of snakes and lizards.

Classification

Desert iguana from Amboy Crater, Mojave Desert, California

Historically, the order Squamata has been divided into three suborders:

• Lacertilia, the lizards
• Serpentes, the snakes (see also Ophidia)
• Amphisbaenia, the worm lizards

Of these, the lizards form a paraphyletic group, since "lizards" excludes the subclades of snakes and amphisbaenians. Studies of squamate relationships using molecular biology have found several distinct lineages, though the specific details of their interrelationships vary from one study to the next. One example of a modern classification of the squamates found the following relationships:

Squamata	Dibamidae Gekkota unnamed Scincomorpha unnamed Lacertoidea (incl. Amphisbaenia) Toxicofera Ophidia (Serpentes) Anguimorpha Iguania

All recent molecular studies suggest that several groups form a venom clade, which encompasses a majority (nearly 60%) of squamate species. Named Toxicofera, it combines the groups Serpentes (snakes), Iguania (agamids, chameleons, iguanids, etc.), and Anguimorpha (monitor lizards, Gila monster, glass lizards, etc.).

List of extant families

Amphisbaenia
Family	Common names	Example species	Example photo
Amphisbaenidae Gray, 1865	Tropical worm lizards	Darwin's worm lizard (Amphisbaena darwinii)	-
Bipedidae Taylor, 1951	Bipes worm lizards	Mexican mole lizard (Bipes biporus)
Blanidae	Mediterranean worm lizards	Mediterranean worm lizard (Blanus cinereus)	-
Cadeidae Vidal & Hedges, 2008	Cuban worm lizards	Cadea blanoides	-
Rhineuridae Vanzolini, 1951	North American worm lizards	North American worm lizard (Rhineura floridana)
Trogonophidae Gray, 1865	Palearctic worm lizards	Checkerboard worm lizard (Trogonophis wiegmanni)	-
Gekkota (incl. Dibamia)
Family	Common names	Example species	Example photo
Dibamidae Boulenger, 1884	Blind lizards	Dibamus nicobaricum	-
Gekkonidae Gray, 1825 (paraphyletic)	Geckos	Thick-tailed gecko (Underwoodisaurus milii)
Pygopodidae Boulenger, 1884	Legless lizards	Burton's snake lizard (Lialis burtonis)
Iguania
Family	Common names	Example species	Example photo
Agamidae Spix, 1825	Agamas	Eastern bearded dragon (Pogona barbata)
Chamaeleonidae Gray, 1825	Chameleons	Veiled chameleon (Chamaeleo calyptratus)
Corytophanidae Frost & Etheridge, 1989	Casquehead lizards	Plumed basilisk (Basiliscus plumifrons)
Crotaphytidae Frost & Etheridge, 1989	Collared and leopard lizards	Common collared lizard (Crotaphytus collaris)
Hoplocercidae Frost & Etheridge, 1989	Wood lizards or clubtails	Club-tail iguana (Hoplocercus spinosus)	-
Iguanidae	Iguanas	Marine iguana (Amblyrhynchus cristatus)
Leiosauridae Frost et al., 2001	-	Darwin's iguana (Diplolaemus darwinii)	-
Liolaemidae Frost & Etheridge, 1989	Swifts	Shining tree iguana (Liolaemus nitidus)
Opluridae Frost & Etheridge, 1989	Madagascan iguanas	Chalarodon (Chalarodon madagascariensis)	-
Phrynosomatidae Frost & Etheridge, 1989	Earless, spiny, tree, side-blotched and horned lizards	Greater earless lizard (Cophosaurus texanus)
Polychrotidae Frost & Etheridge, 1989 (+ Dactyloidae)	Anoles	Carolina anole (Anolis carolinensis)
Tropiduridae Frost & Etheridge, 1989	Neotropical ground lizards	(Microlophus peruvianus)
Lacertoidea (excl. Amphisbaenia)
Family	Common Names	Example Species	Example Photo
Gymnophthalmidae	Spectacled lizards	Bachia bicolor
Lacertidae Oppel, 1811	Wall or true lizards	Ocellated lizard (Lacerta lepida)
Teiidae	Tegus or whiptails	Gold tegu (Tupinambis teguixin)
Neoanguimorpha
Family	Common names	Example species	Example photo
Anguidae Oppel, 1811	Glass lizards, alligator lizards and slow worms	Slow worm (Anguis fragilis)
Anniellidae Gray, 1852	American legless lizards	California legless lizard (Anniella pulchra)
Helodermatidae	Gila monsters	Gila monster (Heloderma suspectum)
Xenosauridae Cope, 1866	Knob-scaled lizards	Mexican knob-scaled lizard (Xenosaurus grandis)
Paleoanguimorpha or Varanoidea
Family	Common names	Example species	Example photo
Lanthanotidae	Earless monitor	Earless monitor (Lanthanotus borneensis)	-
Shinisauridae	Chinese crocodile lizard	Chinese crocodile lizard (Shinisaurus crocodilurus)
Varanidae	Monitor lizards	Perentie (Varanus giganteus)
Scincoidea
Family	Common Names	Example Species	Example Photo
Cordylidae	Spinytail lizards	Girdle-tailed lizard (Cordylus warreni)
Gerrhosauridae	Plated lizards	Sudan plated lizard (Gerrhosaurus major)
Scincidae Oppel, 1811	Skinks	Western blue-tongued skink (Tiliqua occipitalis)
Xantusiidae	Night lizards	Granite night lizard (Xantusia henshawi)
Alethinophidia
Family	Common names	Example species	Example photo
Acrochordidae Bonaparte, 1831	File snakes	Marine file snake (Acrochordus granulatus)
Aniliidae Stejneger, 1907	Coral pipe snakes	Burrowing false coral (Anilius scytale)
Anomochilidae Cundall, Wallach and Rossman, 1993.	Dwarf pipe snakes	Leonard's pipe snake, (Anomochilus leonardi)
Boidae Gray, 1825 (incl. Calabariidae)	Boas	Amazon tree boa (Corallus hortulanus)
Bolyeriidae Hoffstetter, 1946	Round Island boas	Round Island burrowing boa (Bolyeria multocarinata)
Colubridae Oppel, 1811 sensu lato (incl. Dipsadidae, Natricidae, Pseudoxenodontidae)	Colubrids	Grass snake (Natrix natrix)
Cylindrophiidae Fitzinger, 1843	Asian pipe snakes	Red-tailed pipe snake (Cylindrophis ruffus)
Elapidae Boie, 1827	Cobras, coral snakes, mambas, kraits, sea snakes, sea kraits, Australian elapids	King cobra (Ophiophagus hannah)
Homalopsidae Bonaparte, 1845			-
Lamprophiidae Fitzinger, 1843		Bibron's burrowing asp (Atractaspis bibroni)
Loxocemidae Cope, 1861	Mexican burrowing snakes	Mexican burrowing snake (Loxocemus bicolor)
Pareatidae Romer, 1956			-
Pythonidae Fitzinger, 1826	Pythons	Ball python (Python regius)
Tropidophiidae Brongersma, 1951	Dwarf boas	Northern eyelash boa (Trachyboa boulengeri)
Uropeltidae Müller, 1832	Shield-tailed snakes, short-tailed snakes	Cuvier's shieldtail (Uropeltis ceylanica)
Viperidae Oppel, 1811	Vipers, pitvipers, rattlesnakes	European asp (Vipera aspis)
Xenodermatidae Fitzinger, 1826			-
Xenopeltidae Gray, 1849	Sunbeam snakes	Sunbeam snake (Xenopeltis unicolor)
Scolecophidia (incl. Anomalepidae)
Family	Common names	Example species	Example photo
Anomalepidae Taylor, 1939	Dawn blind snakes	Dawn blind snake (Liotyphlops beui)
Gerrhopilidae Vidal et al., 2010			-
Leptotyphlopidae Stejneger, 1892	Slender blind snakes	Texas blind snake (Leptotyphlops dulcis)
Typhlopidae Merrem, 1820	Blind snakes	European blind snake (Typhlops vermicularis)
Xenotyphlopidae Vidal et al., 2010		Xenotyphlops grandidieri	-

Further reading

• Evans SE. 2008. The skull of lizards and tuatara. In Biology of the Reptilia, Vol.20, Morphology H: the skull of Lepidosauria, Gans C, Gaunt A S, Adler K. (eds). Ithica, New York, Society for the study of Amphibians and Reptiles. pp1–344. Weblink to purchase

External links

• Squamata