Squamata

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Squamates
Temporal range: Bathonian–Present [1]
Squamata-01.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Superorder: Lepidosauria
Order: Squamata
Oppel, 1811
Subgroups [2]

Squamata ( /skwæˈmtə/ , Latin squamatus, 'scaly, having scales') is the largest order of reptiles, comprising lizards (including snakes). With over 12,162 species, [3] 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 Diet. 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).

Contents

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 clade Lepidosauria.

Evolution

The holotype of Slavoia darevskii, a fossil squamate Slavoia darevskii.jpg
The holotype of Slavoia darevskii , a fossil squamate

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 subclass 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. [4] [5]

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. [6] [7] However, a 2021 study found the genus to be a lepidosaur of uncertain position, in a polytomy with Squamata and Rhynchocephalia. [8]

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. [9] A 2023 study found that Cryptovaranoides most likely represents an archosauromorph with no apparent squamate affinities, [10] though the original describers maintained their original conclusion that this taxon represents a squamate. [11] The oldest unambiguous fossils of Squamata date to the Bathonian age of the Middle Jurassic of the Northern Hemisphere, [1] with the first appearance of many modern groups, including snakes, during this period. [12]

Scientists believe crown group squamates probably originated in the Early Jurassic based on the fossil record, [4] with the oldest unambiguous fossils of squamates dating to the Middle Jurassic. [1] Squamate morphological and ecological diversity substantially increased over the course of the Cretaceous, [12] 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. [13] Squamates suffered a mass extinction at the Cretaceous–Paleogene (K–Pg) boundary, which wiped out polyglyphanodontians, mosasaurs, and many other distinct lineages. [14]

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, [13] but genetic data suggest that geckos are the earliest crown group squamates. [15] 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.

Reproduction

Trachylepis maculilabris skinks mating Trachylepis maculilabris mating.jpg
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 mammalian penis. [16] 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. [17] [18] Some species, such as the Komodo dragon, can reproduce asexually through parthenogenesis. [19]

The Japanese striped snake has been studied in sexual selection. Elaphe quadrivirgata.JPG
The Japanese striped snake has been studied in sexual selection.

Studies have been conducted on how sexual selection manifests itself in snakes and lizards. Snakes use a variety of tactics in acquiring mates. [20] [ dubious discuss ] Ritual combat between males for the females with which they want to mate includes topping, a behavior exhibited by most viperids, in which one male twists around the vertically elevated fore body of his opponent and forcing it downward. Neck biting commonly occurs while the snakes are entwined. [21]

Facultative parthenogenesis

The effects of central fusion and terminal fusion on heterozygosity Central fusion and terminal fusion automixis.svg
The effects of central fusion and terminal fusion on heterozygosity

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 (cottonmouth snake) can reproduce by facultative parthenogenesis; they are capable of switching from a sexual mode of reproduction to an asexual mode. [22] 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. [22]

Reproduction in squamate reptiles is ordinarily sexual, with males having a ZZ pair of sex-determining chromosomes, and females a ZW pair. However, the Colombian rainbow boa, Epicrates maurus , can also reproduce by facultative parthenogenesis, resulting in production of WW female progeny. [23] The WW females are likely produced by terminal automixis.

Inbreeding avoidance

When female sand lizards mate with two or more males, sperm competition within the female's reproductive tract may occur. Active selection of sperm by females appears to occur in a manner that enhances female fitness. [24] On the basis of this selective process, the sperm of males that are more distantly related to the female are preferentially used for fertilization, rather than the sperm of close relatives. [24] This preference may enhance the fitness of progeny by reducing inbreeding depression.

Evolution of 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. [25] The fossil record shows the divergence between anguimorphs, iguanians, and advanced snakes dates back roughly 200 million years ago (Mya) to the Late Triassic/Early Jurassic, [25] but the only good fossil evidence is from the Middle Jurassic. [26]

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. [27] Previous literature hypothesized that venoms were modifications of salivary or pancreatic proteins, [28] but different toxins have been found to have been recruited from numerous different protein bodies and are as diverse as their functions. [29]

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, [30] which leads to a diversification of toxins that allows the ambush predators the ability to attack a wide range of prey. [31] 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. [32]

Humans and squamates

Bites and fatalities

Map showing the global distribution of venomous snakebites Number of snake envenomings (2007).svg
Map showing the global distribution of venomous snakebites

An estimated 125,000 people a year die from venomous snake bites. [33] In the US alone, more than 8,000 venomous snake bites are reported each year, but only one in 50 million people (five or six fatalities per year in the USA) will die from venomous snake bites. [34] [35]

Lizard bites, unlike venomous snake bites, are usually 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. [36] The Gila monster and beaded lizards of North and Central America are venomous, but not deadly to humans.

Conservation

Though they survived the Cretaceous–Paleogene extinction event, many squamate species are now endangered 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 other anthropogenic causes. Because of this, some squamate species have recently become extinct, with Africa having the most extinct species. Breeding programs and wildlife parks, though, are trying to save many endangered reptiles from extinction. Zoos, private hobbyists, and breeders help educate people about the importance of snakes and lizards.

Classification and phylogeny

Desert iguana from Amboy Crater, Mojave Desert, California DesertIguana031611.jpg
Desert iguana from Amboy Crater, Mojave Desert, California

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

Of these, the lizards form a paraphyletic group, [37] since the "lizards" are found in several distinct lineages, with snakes and amphisbaenians recovered as monophyletic groups nested within. Although studies of squamate relationships using molecular biology have found different relationships between some squamata lineagaes, all recent molecular studies [25] suggest that the venomous groups are united in a venom clade. Named Toxicofera, it encompasses a majority (nearly 60%) of squamate species and includes Serpentes (snakes), Iguania (agamids, chameleons, iguanids, etc.), and Anguimorpha (monitor lizards, Gila monster, glass lizards, etc.). [25]

One example of a modern classification of the squamates is shown below. [2] [38]

Squamata
Dibamia

Dibamidae

Bifurcata
Gekkota
Unidentata
Scinciformata
Episquamata
Laterata
Teiformata

Gymnophthalmidae Merrem 1820 PZSL1851PlateReptilia06 Cercosaura ocellata.png

Teiidae Gray 1827 Bilder-Atlas zur wissenschaftlich-popularen Naturgeschichte der Wirbelthiere (Tupinambis teguixin).jpg

Lacertibaenia
Lacertiformata

Lacertidae Brockhaus' Konversations-Lexikon (1892) (Lacerta agilis).jpg

Amphisbaenia

Rhineuridae Vanzolini 1951

Bipedidae Taylor 1951 Bilder-Atlas zur wissenschaftlich-popularen Naturgeschichte der Wirbelthiere (Bipes canaliculatus).jpg

Blanidae Kearney & Stuart 2004 Blanus cinereus flipped.jpg

Cadeidae Vidal & Hedges 2008

Trogonophidae Gray 1865

Amphisbaenidae Gray 1865 Amphisbaena microcephalum 1847 - white background.jpg

Toxicofera
Serpentes

List of extant families

The over 10,900 extant squamates are divided into 67 families.

Amphisbaenia
FamilyCommon namesExample speciesExample photo
Amphisbaenidae
Gray, 1865
Tropical worm lizardsDarwin's worm lizard ( Amphisbaena darwinii ) Amphisbaenidae - Amphisbaena darwinii.JPG
Bipedidae
Taylor, 1951
Bipes worm lizards Mexican mole lizard (Bipes biporus) Bipes biporus.jpg
Blanidae
Kearney, 2003
Mediterranean worm lizardsMediterranean worm lizard ( Blanus cinereus ) Culebra Ciega - panoramio.jpg
Cadeidae
Vidal & Hedges, 2007 [39]
Cuban worm lizards Cadea blanoides Cadea palirostrata Dickerson 1916.jpg
Rhineuridae
Vanzolini, 1951
North American worm lizards North American worm lizard (Rhineura floridana) Amphisbaenia 1.jpg
Trogonophidae
Gray, 1865
Palearctic worm lizards Checkerboard worm lizard (Trogonophis wiegmanni) Trogonophis wiegmanni imported from iNaturalist photo 24355639 on 14 January 2020.jpg
Gekkota (geckos, incl. Dibamia)
FamilyCommon namesExample speciesExample photo
Carphodactylidae
Kluge, 1967
Southern padless geckos Thick-tailed gecko (Underwoodisaurus milii) Thick-tailed Gecko (Underwoodisaurus milii) (8636512143).jpg
Dibamidae
Boulenger, 1884
Blind lizards Dibamus nicobaricum Anelytropsis.jpg
Diplodactylidae
Underwood, 1954
Australasian geckos Golden-tailed gecko (Strophurus taenicauda) Golden Tailed Gecko.jpg
Eublepharidae
Boulenger, 1883
Eyelid geckos Common leopard gecko (Eublepharis macularius) Eublepharis macularius1.jpg
Gekkonidae
Gray, 1825
Geckos Madagascar giant day gecko (Phelsuma grandis) Madagascar giant day gecko (Phelsuma grandis) Nosy Komba.jpg
Phyllodactylidae
Gamble et al., 2008
Leaf finger geckos Moorish gecko (Tarentola mauritanica) Konstantinos Kalaentzis Tarentola mauritanica (A1).jpg
Pygopodidae
Boulenger, 1884
Flap-footed lizards Burton's snake lizard (Lialis burtonis) Lialis burtonis.jpg
Sphaerodactylidae
Underwood, 1954
Round finger geckos Fantastic least gecko (Sphaerodactylus fantasticus) Sphaerodactylus fantasticus fantasticus (51113243252).jpg
Iguania
FamilyCommon namesExample speciesExample photo
Agamidae
Gray, 1827
Agamas Eastern bearded dragon (Pogona barbata) Bearded dragon04.jpg
Chamaeleonidae
Rafinesque, 1815
Chameleons Veiled chameleon (Chamaeleo calyptratus) Chamaelio calyptratus.jpg
Corytophanidae
Fitzinger, 1843
Casquehead lizards Plumed basilisk (Basiliscus plumifrons) Plumedbasiliskcele4 edit.jpg
Crotaphytidae
H.M. Smith & Brodie, 1982
Collared and leopard lizards Common collared lizard (Crotaphytus collaris) Collared lizard in Zion National Park.jpg
Dactyloidae
Fitzinger, 1843
Anoles Carolina anole (Anolis carolinensis) Anolis carolinensis.jpg
Hoplocercidae
Frost & Etheridge, 1989
Wood lizards or clubtails Enyalioides binzayedi Holotype of Enyalioides binzayedi - ZooKeys-277-069-g007-top.jpg
Iguanidae
Oppel, 1811
Iguanas Marine iguana (Amblyrhynchus cristatus) Marineiguana03.jpg
Leiocephalidae
Frost & Etheridge, 1989
Curly-tailed lizards Hispaniolan masked curly-tailed lizard (Leiocephalus personatus) Leiocephalus-personatus-maskenleguan.jpg
Leiosauridae
Frost et al., 2001
Leiosaurid lizards Enyalius bilineatus Enyalius bilineatus no Parque Estadual de Caparao por Lucas Rosado (08).jpg
Liolaemidae
Frost & Etheridge, 1989
Tree iguanas, snow swifts Shining tree iguana (Liolaemus nitidus) Atacama lizard1.jpg
Opluridae
Titus & Frost, 1996
Malagasy iguanas Chalarodon madagascariensis Chalarodon madagascariensis male.jpg
Phrynosomatidae
Fitzinger, 1843
Earless, spiny, tree, side-blotched and horned lizards Greater earless lizard (Cophosaurus texanus) Reptile tx usa.jpg
Polychrotidae
Frost & Etheridge, 1989
Bush anoles Brazilian bush anole (Polychrus acutirostris) Polychrus acutirostris.JPG
Tropiduridae
Bell, 1843
Neotropical ground lizards Microlophus peruvianus Mperuvianus.jpg
Lacertoidea (excl. Amphisbaenia)
FamilyCommon NamesExample SpeciesExample Photo
Alopoglossidae
Goicoechea, Frost, De la Riva, Pellegrino, Sites Jr., Rodrigues, & Padial, 2016
Alopoglossid lizards Alopoglossus vallensis Ptychoglossus vallensis.jpg
Gymnophthalmidae
Fitzinger, 1826
Spectacled lizards Bachia bicolor Bachia bicolor.jpg
Lacertidae
Oppel, 1811
Wall lizards Ocellated lizard (Lacerta lepida) Perleidechse-20.jpg
Teiidae
Gray, 1827
Tegus and whiptails Gold tegu (Tupinambis teguixin) Goldteju Tupinambis teguixin.jpg
Anguimorpha
FamilyCommon namesExample speciesExample photo
Anguidae
Gray, 1825
Glass lizards, alligator lizards and slowwormsSlowworm ( Anguis fragilis ) Anguidae.jpg
Anniellidae
Boulenger, 1885
American legless lizards California legless lizard (Anniella pulchra) Anniella pulchra.jpg
Diploglossidae
Bocourt, 1873
galliwasps, legless lizards Jamaican giant galliwasp (Celestus occiduus) Celestus occiduus museum specimen.jpeg -
Helodermatidae
Gray, 1837
Beaded lizards Gila monster (Heloderma suspectum) Gila.monster.arp.jpg -
Lanthanotidae
Steindachner, 1877
Earless monitor Earless monitor (Lanthanotus borneensis) Real Lanthanotus borneensis.jpg
Shinisauridae
Ahl, 1930
Chinese crocodile lizard Chinese crocodile lizard (Shinisaurus crocodilurus) Chin-krokodilschwanzechse-01.jpg
Varanidae
Merrem, 1820
Monitor lizards Perentie (Varanus giganteus) Perentie Lizard Perth Zoo SMC Spet 2005.jpg
Xenosauridae
Cope, 1866
Knob-scaled lizards Mexican knob-scaled lizard (Xenosaurus grandis) Xenosaurus grandis.jpg
Scincoidea
FamilyCommon NamesExample SpeciesExample Photo
Cordylidae
Fitzinger, 1826
Girdled lizards Girdle-tailed lizard (Cordylus warreni) Cordylus breyeri1.jpg
Gerrhosauridae
Fitzinger, 1843
Plated lizards Sudan plated lizard (Gerrhosaurus major) Gerrhosaurus major.jpg
Scincidae
Oppel, 1811
Skinks Western blue-tongued skink (Tiliqua occipitalis) Tiliqua occipitalis.jpg
Xantusiidae
Baird, 1858
Night lizards Granite night lizard (Xantusia henshawi) Xantusia henshawi.jpg
Alethinophidia
FamilyCommon namesExample speciesExample photo
Acrochordidae
Bonaparte, 1831 [40]
File snakes Marine file snake (Acrochordus granulatus) Wart snake 1.jpg
Aniliidae
Stejneger, 1907 [41]
Coral pipe snakes Burrowing false coral (Anilius scytale) False Coral Snake (Anilius scytale) close-up (13929278050).jpg
Anomochilidae
Cundall, Wallach and Rossman, 1993. [42]
Dwarf pipe snakes Leonard's pipe snake, (Anomochilus leonardi) Anomochilus weberi.jpg
Boidae
Gray, 1825 [40] (incl. Calabariidae)
Boas Amazon tree boa (Corallus hortulanus) Corallushortulanus.png
Bolyeriidae
Hoffstetter, 1946
Round Island boas Round Island burrowing boa (Bolyeria multocarinata) Round Island Boa.jpeg
Colubridae
Oppel, 1811 [40] sensu lato (incl. Dipsadidae, Natricidae, Pseudoxenodontidae)
Colubrids Grass snake (Natrix natrix) Natrix natrix (Marek Szczepanek).jpg
Cylindrophiidae
Fitzinger, 1843
Asian pipe snakes Red-tailed pipe snake (Cylindrophis ruffus) Cylindrophis rufus.jpg
Elapidae
Boie, 1827 [40]
Cobras, coral snakes, mambas, kraits, sea snakes, sea kraits, Australian elapids King cobra (Ophiophagus hannah) Ophiophagus hannah2.jpg
Homalopsidae
Bonaparte, 1845
Indo-Australian water snakes, mudsnakes, bockadams New Guinea bockadam (Cerberus rynchops) HerpetonTentaculatumFord.jpg
Lamprophiidae
Fitzinger, 1843 [43]
Lamprophiid snakes Bibron's burrowing asp (Atractaspis bibroni) Lamprophis fuliginosus02.jpg
Loxocemidae
Cope, 1861
Mexican burrowing snakes Mexican burrowing snake (Loxocemus bicolor) Loxocemus bicolor.jpg
Pareidae
Romer, 1956
Pareid snakes Perrotet's mountain snake (Xylophis perroteti) Xylophis sp. Munnar.jpg
Pythonidae
Fitzinger, 1826
Pythons Ball python (Python regius) Ball python lucy.JPG
Tropidophiidae
Brongersma, 1951
Dwarf boas Northern eyelash boa (Trachyboa boulengeri) Cuban Giant Trope (Tropidophis melanurus) (8577519420).jpg
Uropeltidae
Müller, 1832
Shield-tailed snakes, short-tailed snakes Cuvier's shieldtail (Uropeltis ceylanica) Silybura shortii.jpg
Viperidae
Oppel, 1811 [40]
Vipers, pitvipers, rattlesnakes European asp (Vipera aspis) Vipera aspis aspis, Lorraine, France.jpg
Xenodermidae
Fitzinger, 1826
Odd-scaled snakes and relatives Khase earth snake (Stoliczkia khasiensis) Achalinus formosanus formosanus full body shot.jpg
Xenopeltidae
Gray, 1849
Sunbeam snakes Sunbeam snake (Xenopeltis unicolor) XenopeltisUnicolorRooij.jpg
Scolecophidia (incl. Anomalepidae)
FamilyCommon namesExample speciesExample photo
Anomalepidae
Taylor, 1939 [40]
Dawn blind snakes Dawn blind snake (Liotyphlops beui)
Gerrhopilidae
Vidal et al., 2010 [39]
Indo-Malayan blindsnakes Andaman worm snake (Gerrhopilus andamanensis)
Leptotyphlopidae
Stejneger, 1892 [40]
Slender blind snakes Texas blind snake (Leptotyphlops dulcis) Leptotyphlops dulcis.jpg
Typhlopidae
Merrem, 1820 [44]
Blind snakes European blind snake (Typhlops vermicularis) Typhlops vermicularis.jpg
Xenotyphlopidae
Vidal et al., 2010 [39]
Malagasy blind snakes Xenotyphlops grandidieri

References

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Further reading