Amphisbaenia (called amphisbaenians or worm lizards) is a group of usually legless

squamate Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest order of reptiles, comprising lizards, snakes, and amphisbaenians (worm lizards), which are collectively known as squamates or scaled reptiles. With over 10,900 species, it ...

s, comprising over 200 extant species. Amphisbaenians are characterized by their long bodies, the reduction or loss of the limbs, and rudimentary eyes. As many species have a pink body and scales arranged in rings, they have a superficial resemblance to earthworms. While the genus '' Bipes'' retains forelimbs, all other genera are limbless. Although superficially similar to the

snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more j ...

s and blind lizards, recent phylogenetic studies suggest that they are most closely related to wall lizards of the family Lacertidae. Amphisbaenians are widely distributed, occurring in North America, Europe, Africa, South America, Western Asia and the Caribbean. Most species are less than long.

Description

200px, left, Close-up of the head of'' Rhineura'' Despite a superficial resemblance to some primitive snakes, amphisbaenians have many unique features that distinguish them from other reptiles. Internally, their right lung is reduced in size to fit their narrow bodies, whereas in snakes, it is always the left lung. Their skeletal structure and skin are also different from those of other squamates. Both genetic and recent fossil evidence indicate that amphisbaenians lost their legs independently from snakes. The head is stout, not set off from the neck, and either rounded, sloped, or sloped with a ridge down the middle. Most of the skull is solid bone, with a distinctive single median tooth in the upper jaw. It has no outer ears, and the eyes are deeply recessed and covered with skin and scales. These rudimentary eyes have a

cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...

lens A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements ...

, and complex

ciliary body The ciliary body is a part of the eye that includes the ciliary muscle, which controls the shape of the lens, and the ciliary epithelium, which produces the aqueous humor. The aqueous humor is produced in the non-pigmented portion of the ciliar ...

, which allows them to detect light, but they are reduced in size and do not have an

anterior chamber The anterior chamber ( AC) is the aqueous humor-filled space inside the eye between the iris and the cornea's innermost surface, the endothelium. Hyphema, anterior uveitis and glaucoma are three main pathologies in this area. In hyphema, blood f ...

. The body is elongated, and the tail truncates in a manner that vaguely resembles the head. Their name is derived from Amphisbaena, a mythical serpent with a head at each end—referencing both the manner in which their tail truncates, and their ability to move just as well in reverse as forwards. The four species of '' Bipes'' are unusual in having a pair of forelimbs, but all limbless species have some remnants of the pelvic and pectoral girdles embedded within the body musculature. Blanus skull by Nick Longrich

Amphisbaenians have a distinctive skin made up of rings of scales (annuli) that form a tube in which the loosely attached trunk of the body moves. Burrowing is achieved with an accordion-like motion, with longitudinal muscles in the skin bunching up the annuli, anchoring it to the surrounding soil, and trunk muscles moving the body forward or backwards within the integumentary tube. Amphisbaenians are carnivorous, able to tear chunks out of larger prey with their powerful, interlocking teeth. Like lizards, some species are able to shed their tails (

autotomy Autotomy (from the Greek ''auto-'', "self-" and ''tome'', "severing", αὐτοτομία) or self-amputation, is the behaviour whereby an animal sheds or discards one or more of its own appendages, usually as a self-defense mechanism to elude ...

). Most species lay eggs, although at least some are known to be

viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the ...

. The white worm lizard (''Amphisbaena alba'') is often found in association with

leafcutter ant Leafcutter ants, a non-generic name, are any of 47 species of leaf-chewing ants belonging to the two genera '' Atta'' and ''Acromyrmex''. These species of tropical, fungus-growing ants are all endemic to South and Central America, Mexico, and par ...

s. This reptile is thought to forage in the ants' deep galleries, where the insects deposit their waste. The presence of these reptiles is easily explained by the fact that they prey on the larvae of large beetles that also inhabit the leafcutter ants' galleries. Amphisbaenians have often been categorized by their skull shape. The specialized skull shape is hypothesized to be driven by environmental and ecological conditions, such as soil type, and is an instance of

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...

Distribution

Amphisbaenians are found in North America, Europe, Africa, South America, the Middle East, and the Caribbean, a surprisingly large distribution despite being small subterranean animals that rarely ever leave their burrows. Initially, this large distribution was thought to be due to

vicariance Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...

, or the result of the breakup of Pangaea. This hypothesis was supported by morphological data that dated amphisbaenian diversification to over 200 million years ago (Mya), while Pangaea was still intact. However, a recent study using a combination of molecular and fossil evidence suggests that amphisbaenians originated in North America, where they underwent their first divergence around 107 Mya.Longrich, N., Vinter, J., Pisani, D., Pyron, A., Gauthier, J. 2015 Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. ''Proceedings of the Royal Society B'', 282 http://rspb.royalsocietypublishing.org/content/282/1806/20143034 They then underwent another major diversification into North American and European forms 40–56 Mya. Finally, the African and South American forms split around 40 Mya. This suggests that worm-lizards crossed the Atlantic Ocean (which had fully formed by 100 Mya) twice, once just after the K–Pg extinction, and then again, later in the

Palaeogene The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago ( Mya) to the beginning o ...

. This also implies that limblessness evolved independently three times, a finding that contrasts the morphological theory that limbed amphisbaenians are the most basal. This widespread dispersal is suggested to have occurred by rafting – natural erosion or a storm event loosened a large raft of soil and vegetation that drifted across the ocean until landing on another shore. This oceanic rafting would be feasible due to the subterranean lifestyle and small nutritional requirements of amphisbaenids. After the

Chicxulub impact The Chicxulub crater () is an impact crater buried underneath the Yucatán Peninsula in Mexico. Its center is offshore near the community of Chicxulub, after which it is named. It was formed slightly over 66 million years ago when a large a ...

, many predators of amphisbaenians became extinct, which allowed colonist amphisbaenians to thrive in new territories.

Evolution

The oldest

stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

amphisbaenian, the limbed ''Slavoia darevskii'' is known from the Late Cretaceous (

Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...

) of Mongolia. The oldest known modern amphisbaenians are members of

Rhineuridae Rhineuridae is a family of amphisbaenians (commonly called worm lizards) that includes one living genus and species, ''Rhineura floridana'', as well as many extinct species belonging to both ''Rhineura'' and several extinct genera. The living ''R ...

and the extinct family Oligodontosauridae from the

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...

of North America. Modern amphisbaenians likely originated in North America, before dispersing to South America, Africa and Europe via

rafting Rafting and whitewater rafting are recreational outdoor activities which use an inflatable raft to navigate a river or other body of water. This is often done on whitewater or different degrees of rough water. Dealing with risk is often a ...

during the Paleogene.

Taxonomy

Taxonomic classification of amphisbaenians was traditionally based on morphological characters, such as the number of preanal pores, body annuli, tail annuli, and skull shape. Such characters are vulnerable to convergent evolution; in particular, the loss of the forelimbs and the evolution of specialized shovel-headed and keel-headed morphs appear to have occurred multiple times in the history of the group. Classifications based on mitochondrial DNA sequences and

nuclear DNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. I ...

sequences better reflect their true evolutionary history, and are now being used to distinguish genera of amphisbaenians. The most ancient branch of the tree is the Rhineuridae. The remaining five families form a group to the exclusion of rhineurids. Bipedidae, Blanidae, and Cadeidae represent the most ancient divergences within this grouping, with Trogonophidae and Amphisbaenidae diverging more recently. South American amphisbaenids apparently are derived from African amphisbaenids that rafted across the Atlantic in the

Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...

, about 40 million years ago. Cuban cadeids may be similarly derived from blanids that rafted across from northwestern Africa or southwestern Europe in a similar time frame. Amphisbaenia has usually been considered a suborder of squamates, but more recent studies indicate that it is part of the lizard clade

Lacertoidea The Lacertoidea is a group of squamate reptiles that includes the Lacertidae, Teiidae, Gymnophthalmidae, and Amphisbaenia. The finding from molecular phylogenetic studies that the burrowing Amphisbaenia were nested in a clade with the lizard f ...

, ranked only as a superfamily, so it is now commonly described as an unranked clade.

Families

Six families of amphisbaenians are currently recognised: *

Amphisbaenidae The Amphisbaenidae (common name: worm lizards) are a family of amphisbaenians, a group of limbless vertebrates. Geographic range Amphisbaenids occur in South America, some Caribbean islands, Europe, and sub-Saharan Africa. Taxonomy One deep- ...

Gray Grey (more common in British English) or gray (more common in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning literally that it is "without color", because it can be composed o ...

, 1865 – Amphisbaenids, tropical worm lizards of South America, some Caribbean islands, and Sub-Saharan Africa. (12 genera, 182 species) *

Bipedidae Bipedidae is a family of amphisbaenians that includes the extant genus '' Bipes'' represented by three species from Baja California and the southern coast of Mexico and the extinct genus '' Anniealexandria'' represented by one species that lived ...

Taylor Taylor, Taylors or Taylor's may refer to: People * Taylor (surname) ** List of people with surname Taylor * Taylor (given name), including Tayla and Taylah * Taylor sept, a branch of Scottish clan Cameron * Justice Taylor (disambiguation) Pl ...

, 1951 – Only in Mexico and commonly called ''ajolotes'', but not to be confused with

axolotl The axolotl (; from nci, āxōlōtl ), ''Ambystoma mexicanum'', is a paedomorphic salamander closely related to the tiger salamander. Axolotls are unusual among amphibians in that they reach adulthood without undergoing metamorphosis. I ...

s (1 genus, 3 species) *

Blanidae ''Blanus'' is a genus of amphisbaenians found in the Mediterranean region of Europe and North Africa. Like other amphisbaenians, ''Blanus'' species are specialized for a subterranean existence, with long, slender bodies, reduced limbs, and rudim ...

Kearney & Stuart, 2004 – Anatolian, Iberian, and Moroccan worm lizards (1 genus, 7 species) * Cadeidae Vidal and Hedges, 2008 – Cuban keel-headed worm lizards (1 genus, 2 species). Traditionally amphisbaenids, but shown by DNA to be closest to Blanidae. *

Vanzolini, 1951 – North American worm lizards (1 genus) * Trogonophidae Gray, 1865 – Palearctic worm lizards (4 genera, 6 species)

Phylogeny

The following cladogram shows the relationships between the six amphisbaenian families determined in the phylogenetic analysis of mitochondrial and nuclear genes by Vidal et al. (2008).