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The Scolecophidia, commonly known as blind snakes or thread snakes, are an
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
of
snakes Snakes are elongated Limbless vertebrate, limbless reptiles of the suborder Serpentes (). Cladistically Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales much like other members of ...
. They range in length from . All are
fossorial A fossorial animal () is one that is adapted to digging and which lives primarily (but not solely) underground. Examples of fossorial vertebrates are Mole (animal), moles, badgers, naked mole-rats, meerkats, armadillos, wombats, and mole salamand ...
(adapted for burrowing). Five
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
and 39
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
are recognized. The Scolecophidia infraorder is most-likely
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
(with the family
Anomalepididae The Anomalepididae are a Family (biology), family of nonvenomous snakes, native to Central and South America. They are similar to Typhlopidae, except that some species possess a single tooth in the lower jaw. Currently, four Genus, genera and 15 ...
recovered with strong support as sister clade to the 'typical snakes').


Taxonomy

The infraorder name Scolecophidia derives from the two
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
words or σκώληκος (, genitive ), meaning "earthworm", and (), meaning "snake". It refers to their shape and fossorial lifestyle.


Families


Evolution

Despite only having fossils as early as the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
, Scolecophidia itself likely originated in the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
, with
Anomalepididae The Anomalepididae are a Family (biology), family of nonvenomous snakes, native to Central and South America. They are similar to Typhlopidae, except that some species possess a single tooth in the lower jaw. Currently, four Genus, genera and 15 ...
, Leptotyphlopidae, and Typhlopoidea diverging from one another during the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
. Within Typhlopoidea, Gerrhopilidae likely diverged from the Xenotyphlopidae-Typhlopidae
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
during the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
, and
Xenotyphlopidae ''Xenotyphlops'' is a genus of snakes, the only genus of the Family (biology), family Xenotyphlopidae, comprising two species found only in Madagascar. These snakes are also known as the Malagasy blind snake. Evolution ''Xenotyphlops'' is an anc ...
and
Typhlopidae The Typhlopidae are a family of blind snakes. They are found mostly in the tropical regions of Africa, Asia, the Americas, and all mainland Australia and various islands. The rostral scale overhangs the mouth to form a shovel-like burrowing str ...
likely diverged from one another during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
. Scolecophidians are believed to have originated on
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
, with anomalepidids and leptotyphlopids evolving in west Gondwana (South America and Africa) and the Typhlopoidea (typhlopids, gerrhopilids, and xenotyphlopids) on east Gondwana, initially on the combined
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
/
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
land mass, during the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
. Typhlopids, initially isolated on Madagascar, then dispersed to Africa and Eurasia. South American typhlopids appear to have evolved from African typhlopids that rafted across the Atlantic about 60 million years ago; they, in turn, dispersed to the Caribbean about 33 million years ago. Similarly, typhlopids appear to have reached Australia from Southeast Asia or Indonesia about 28 million years ago. Meanwhile, the gerrhopilids, isolated on Insular India, underwent a radiation throughout tropical Asia following the collision of India with Asia, while the xenotyphlopids remained isolated on Madagascar. The Malagasy typhlopoids ('' Madatyphlops'' in Typhlopidae and ''
Xenotyphlops ''Xenotyphlops'' is a genus of snakes, the only genus of the Family (biology), family Xenotyphlopidae, comprising two species found only in Madagascar. These snakes are also known as the Malagasy blind snake. Evolution ''Xenotyphlops'' is an anc ...
'' in Xenotyphlopidae) are among the only extant terrestrial vertebrates on Madagascar whose isolation occurred 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 ...
from the Cretaceous breakup of Gondwana. The only other terrestrial vertebrate on Madagascar that shares this evolutionary history is the Madagascan big-headed turtle (''Erymnochelys madagascariensis''); all other Malagasy land vertebrates dispersed from the mainland to an already-isolated Madagascar from the latest Cretaceous to the present.


Fossil record

The extinct fossil species '' Boipeba tayasuensis'' from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
of
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
was described in 2020, marking the earliest fossil record of Scolecophidia. It was a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to Typhlopoidea and was over 1 meter in length, making it much larger than most modern blindsnakes, with only ''
Afrotyphlops schlegelii ''Afrotyphlops schlegelii'', commonly known as Schlegel's beaked blind snake:fr:William Roy Branch, Branch, Bill (2004). ''Field Guide to Snakes and other Reptiles of Southern Africa''. Third Revised edition, Second impression. Sanibel Island, Fl ...
'' and ''
Afrotyphlops mucruso The African giant blind snake (''Afrotyphlops mucruso''), also called the Zambezi beaked blind snake, is a species of snake in the Typhlopidae family.McDiarmid RW, Campbell JA, Touré T. 1999. Snake Species of the World: A Taxonomic and Geograph ...
'' rivaling it in size. Prior to this, the earliest scolecophidian fossils were only known from the
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
of
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
and the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. Possible Typhopid skin has been identified in
Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil in ...
.


Phylogeny

This phylogeny combines the ones recovered by Vidal ''et al.'' in 2010 and Fachini ''et al.'' in 2020.


Description

The common name of Scolecophidia, blind snakes, is based on their shared characteristic of reduced eyes that are located under their head scales. These head scales are found in all snakes and are referred to as spectacles, but within this infraorder, they are opaque, resulting in decreased visual capabilities. Reduced eyes of the Scolecophidia have been attributed to evolutionary origins of snakes, which are hypothesized to have arisen from fossorial ancestors, causing a loss of genes related to eyesight that later evolved again in higher snakes to be similar to other vertebrates due to convergent evolution. Newer research shows that seven of the 12 genes associated with bright-light vision in most snakes and lizards are not present in this infraorder, and the common ancestor of all snakes had better eyesight.Burrowing snakes have far worse eyesight than their ancestors
/ref> Other shared characteristics include an absent left oviduct in four of the five families, aside from the Anomalepididae, which have a well developed yet reduced left oviduct. Aside from this, these snakes range in length from . Their typical body shapes include slender, cylindrical bodies and small, narrow heads. All these families either lack or have a vestigial left lung and lack cranial infrared receptors.


Behavior

The main shared characteristic found across all Scolecophidia is a fossorial nature, either living underground or within logs and leaf litter. Aside from this, thus far the reproduction remains understudied with all Scolecophidia studied thus far being noted to be oviparous, with elongate eggs noted in both leptotyphlopids and typhlopids. Foraging behaviors vary across families, but all feed on invertebrates. Some of their main food sources include ant or termite eggs, which are tracked down by following chemical cues left by these invertebrates to create trails. ''Tricheilostomata macrolepis'' has been seen climbing up trees and waving its head side to side vertically to detect chemical cues in the air to locate insect nests. In a study on the Leptotyphlopidae, some species were found to specialize in eating only termites or ants; some rely on binge feeding patterns, while others do not. While these snakes are often difficult to locate due to their burrowing habits, they are more often seen above ground after rain due to flooding that occurs in burrows. The ancestral nature of the Scolecophidia has resulted in the use of these organisms as models for evolutionary studies in Serpentes to better understand evolution of reproduction, morphology, and feeding habits.


References

{{Taxonbar, from=Q201636 Taxa named by Edward Drinker Cope Extant Coniacian first appearances