Scolecophidia: Difference between revisions

Scolecophidia; Temporal range: 87.8–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Late Cretaceous - present (but see text)
	Brahminy blind snake,; Ramphotyphlops braminus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Squamata
Suborder:	Serpentes
Infraorder:	Scolecophidia; Cope, 1864
Families
	See text

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The Scolecophidia, commonly known as blind snakes or thread snakes,^[2] are an infraorder^[2] of snakes.^[3] They range in length from 10 to 100 centimetres (4 to 40 inches). All are fossorial (adapted for burrowing).^[4] Five families and 39 genera are recognized.^[5] The Scolecophidia infraorder is most likely paraphyletic (with the family Anomalepididae recovered with strong support as sister clade to the 'typical snakes').^[6]

Taxonomy

The infraorder name Scolecophidia derives from the two Ancient Greek words σκώληξ or σκώληκος (skṓlēx, genitive skṓlēkos), meaning "earthworm", and ὄφις (óphis), meaning "snake".^[7]^[8] It refers to their shape and fossorial lifestyle.

Families

Family^[2]	Authority^[2]	Genera^[5]	Common name^[2]	Geographic range^[1]
Anomalepididae	Taylor, 1939	4	primitive blind snakes	Southern Central America and South America
Gerrhopilidae	Vidal, Wynn, Donnellan & Hedges, 2010^[9]	2	Indo-Malayan blind snakes	India, Southeast Asia, Indonesia, the Philippines, and New Guinea^[10]
Leptotyphlopidae	Stejneger, 1892	13	slender blind snakes or threadsnakes	Africa, western Asia, and the Americas
Typhlopidae	Merrem, 1820	18	long-tailed blind snakes	Most tropical and many subtropical regions all over the world
Xenotyphlopidae	Vidal, Vences, Branch & Hedges, 2010^[9]	1	Malagasy blind snakes	Madagascar

Evolution

Despite only having fossils as early as the Cretaceous, Scolecophidia itself likely originated in the Middle Jurassic, with Anomalepididae, Leptotyphlopidae, and Typhlopoidea diverging from one another during the Late Jurassic. Within Typhlopoidea, Gerrhopilidae likely diverged from the Xenotyphlopidae-Typhlopidae clade during the Early Cretaceous, and Xenotyphlopidae and Typhlopidae likely diverged from one another during the Late Cretaceous.^[9]

Scolecophidians are believed to have originated on Gondwana, 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/Madagascar land mass, during the Mesozoic.^[9] 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.^[9] Similarly, typhlopids appear to have reached Australia from Southeast Asia or Indonesia about 28 million years ago.^[9] 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.^[11]

The Malagasy typhlopoids (Madatyphlops in Typhlopidae and Xenotyphlops in Xenotyphlopidae) are among the only extant terrestrial vertebrates on Madagascar whose isolation occurred due to vicariance 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.^[11]

Fossil record

Illustration of *Boipeba*, the earliest known fossil blind snake

The extinct fossil species Boipeba tayasuensis from the Late Cretaceous of Brazil was described in 2020, marking the earliest fossil record of Scolecophidia. It was a sister group to Typhlopoidea and was over 1 meter in length, making it much larger than most modern blindsnakes, with only Afrotyphlops schlegelii and Afrotyphlops mucruso rivaling it in size. Prior to this, the earliest scolecophidian fossils were only known from the Paleocene of Morocco and the Eocene of Europe.^[12]

Possible Typhopid skin has been identified in Dominican amber.^[13]

Phylogeny

This phylogeny combines the ones recovered by Vidal et al. in 2010 and Fachini et al. in 2020.^[9]^[12]

Scolecophidia

†Boipeba

	Xenotyphlopidae

	Typhlopidae

Description

The common name of Scolecophidia, blind snakes, is based on their shared characteristic of reduced eyes that are located under their head scales.^[14] 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.^[3] 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.^[14] 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.^[15] 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.^[3] Aside from this, these snakes range in length from 10 to 100 cm (4 to 39 in). Their typical body shapes include slender, cylindrical bodies and small, narrow heads.^[14] All these families either lack or have a vestigial left lung and lack cranial infrared receptors.^[3]

Behavior

The main shared characteristic found across all Scolecophidia is a fossorial nature, either living underground or within logs and leaf litter.^[3] Aside from this, thus far the reproduction remains understudied with all Scolecophidia studied thus far being noted to be oviparous,^[3] with elongate eggs noted in both leptotyphlopids and typhlopids.^[16] 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.^[16] 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.^[3] 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.^[3] 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.^[16]

References

^ ^a ^b McDiarmid RW, Campbell JA, Touré TA (1999). Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Washington, District of Columbia: Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
^ ^a ^b ^c ^d ^e "Scolecophidia". Integrated Taxonomic Information System. Retrieved 14 August 2007.
^ ^a ^b ^c ^d ^e ^f ^g ^h Vitt, Laurie J.; Caldwell, Janalee P. (2014). Herpetology: An Introductory Biology of Amphibians and Reptiles (Fourth ed.). Academic Press. pp. 597–603. ISBN 978-0-12-386919-7.
^ Scolecophidia at Palaeos. Accessed 21 December 2013.
^ ^a ^b Uetz, Peter. "The Reptile Database". The Reptile Database. Retrieved 31 December 2017.
^ Miralles, Aurélien; Marin, Julie; Markus, Damin; Herrel, Anthony; Hedges, Blair; Vidal, Nicolas (2018). "Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications". Journal of Evolutionary Biology. 31 (12): 1782–1793. doi:10.1111/jeb.13373. PMID 30193402. S2CID 52174313.
^ Bailly, Anatole (1981-01-01). Abrégé du dictionnaire grec français. Paris: Hachette. ISBN 978-2010035289. OCLC 461974285.
^ Bailly, Anatole. "Greek-french dictionary online". www.tabularium.be. Retrieved January 7, 2019.
^ ^a ^b ^c ^d ^e ^f ^g Vidal, Nicolas; et al. (2010). "Blindsnake evolutionary tree reveals long history on Gondwana". Biology Letters. 6 (4): 558–561, page 560. doi:10.1098/rsbl.2010.0220. PMC 2936224. PMID 20356885.
^ Vitt, Laurie J.; Caldwell, Janalee P. (2013). Herpetology: An Introductory Biology of Amphibians and Reptiles. Academic Press. p. 600. ISBN 9780123869203.
^ ^a ^b Ali, Jason R.; Hedges, S. Blair (2023-05-04). "The colonisation of Madagascar by land‐bound vertebrates". Biological Reviews. doi:10.1111/brv.12966. ISSN 1464-7931.
^ ^a ^b Thiago Schineider Fachini; Silvio Onary; Alessandro Palci; Michael S.Y. Lee; Mario Bronzati; Annie Schmaltz Hsiou (2020). "Cretaceous blindsnake from Brazil fills major gap in snake evolution". iScience. 23 (12): Article 101834. doi:10.1016/j.isci.2020.101834. PMC 7718481. PMID 33305189.
^ Poinar, George O.; Poinar, Roberta (1999). The Amber Forest: A Reconstruction of a Vanished World. Princeton University Press. ISBN 978-0-691-05728-6.
^ ^a ^b ^c Simões, B. F.; et al. (2015). "Visual system evolution and the nature of the ancestral snake". Journal of Evolutionary Biology. 28 (7): 1309–1320. doi:10.1111/jeb.12663. PMID 26012745. S2CID 24013194.
^ Burrowing snakes have far worse eyesight than their ancestors
^ ^a ^b ^c Webb, J.K.; Shine, R.; Branch, W.R.; Harlow, P.S. (2000). "Life‐history strategies in basal snakes: reproduction and dietary habits of the African thread snake Leptotyphlops scutifrons (Serpentes: Leptotyphlopidae)". Journal of Zoology. 250 (3): 321–327. doi:10.1017/s0952836900003058.

[McD99-1] McDiarmid RW, Campbell JA, Touré TA (1999). Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Washington, District of Columbia: Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).

[ITIS-2] "Scolecophidia". Integrated Taxonomic Information System. Retrieved 14 August 2007.

[Vitt2014-3] ^ ^a ^b ^c ^d ^e ^f ^g ^h Vitt, Laurie J.; Caldwell, Janalee P. (2014). Herpetology: An Introductory Biology of Amphibians and Reptiles (Fourth ed.). Academic Press. pp. 597–603. ISBN 978-0-12-386919-7.

[4] Scolecophidia at Palaeos. Accessed 21 December 2013.

[TRD-5] Uetz, Peter. "The Reptile Database". The Reptile Database. Retrieved 31 December 2017.

[6] Miralles, Aurélien; Marin, Julie; Markus, Damin; Herrel, Anthony; Hedges, Blair; Vidal, Nicolas (2018). "Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications". Journal of Evolutionary Biology. 31 (12): 1782–1793. doi:10.1111/jeb.13373. PMID 30193402. S2CID 52174313.

[7] Bailly, Anatole (1981-01-01). Abrégé du dictionnaire grec français. Paris: Hachette. ISBN 978-2010035289. OCLC 461974285.

[8] Bailly, Anatole. "Greek-french dictionary online". www.tabularium.be. Retrieved January 7, 2019.

[Vidal-2010-9] ^ ^a ^b ^c ^d ^e ^f ^g Vidal, Nicolas; et al. (2010). "Blindsnake evolutionary tree reveals long history on Gondwana". Biology Letters. 6 (4): 558–561, page 560. doi:10.1098/rsbl.2010.0220. PMC 2936224. PMID 20356885.

[10] Vitt, Laurie J.; Caldwell, Janalee P. (2013). Herpetology: An Introductory Biology of Amphibians and Reptiles. Academic Press. p. 600. ISBN 9780123869203.

[:1-11] Ali, Jason R.; Hedges, S. Blair (2023-05-04). "The colonisation of Madagascar by land‐bound vertebrates". Biological Reviews. doi:10.1111/brv.12966. ISSN 1464-7931.

[:0-12] Thiago Schineider Fachini; Silvio Onary; Alessandro Palci; Michael S.Y. Lee; Mario Bronzati; Annie Schmaltz Hsiou (2020). "Cretaceous blindsnake from Brazil fills major gap in snake evolution". iScience. 23 (12): Article 101834. doi:10.1016/j.isci.2020.101834. PMC 7718481. PMID 33305189.

[13] Poinar, George O.; Poinar, Roberta (1999). The Amber Forest: A Reconstruction of a Vanished World. Princeton University Press. ISBN 978-0-691-05728-6.

[Simões2015-14] Simões, B. F.; et al. (2015). "Visual system evolution and the nature of the ancestral snake". Journal of Evolutionary Biology. 28 (7): 1309–1320. doi:10.1111/jeb.12663. PMID 26012745. S2CID 24013194.

[15] Burrowing snakes have far worse eyesight than their ancestors

[Webb2000-16] Webb, J.K.; Shine, R.; Branch, W.R.; Harlow, P.S. (2000). "Life‐history strategies in basal snakes: reproduction and dietary habits of the African thread snake Leptotyphlops scutifrons (Serpentes: Leptotyphlopidae)". Journal of Zoology. 250 (3): 321–327. doi:10.1017/s0952836900003058.

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@@ Line 3: / Line 3: @@
 | name = Scolecophidia
 | fossil_range = {{Fossil range|87.8|0}} [[Late Cretaceous]] - present (but see text)
-| image = Ramphotyphlops braminus.jpg
+| image = Ramphotyphlops braminus jja.jpg
 | image_caption = Brahminy blind snake,<br />''[[Indotyphlops braminus|Ramphotyphlops braminus]]''
 | taxon = Scolecophidia
-| authority = [[Edward Drinker Cope|Cope]], 1864<ref name="McD99">[[:fr:Roy Wallace McDiarmid|McDiarmid RW]], [[Jonathan A. Campbell|Campbell JA]], Touré TA (1999). ''Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1''. Washington, District of Columbia: Herpetologists' League. 511 pp. {{ISBN|1-893777-00-6}} (series). {{ISBN|1-893777-01-4}} (volume).</ref>
+| authority = [[Edward Drinker Cope|Cope]], 1864<ref name="McD99">[[species:Roy Wallace McDiarmid|McDiarmid RW]], [[Jonathan A. Campbell|Campbell JA]], [[species:T’Shaka A. Touré|Touré TA]] (1999). ''Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1''. Washington, District of Columbia: Herpetologists' League. 511 pp. {{ISBN|1-893777-00-6}} (series). {{ISBN|1-893777-01-4}} (volume).</ref>
 | subdivision_ranks = Families
 | subdivision = See text
 }}
-The '''Scolecophidia''', commonly known as '''blind snakes''' or '''thread snakes''',<ref name="ITIS">{{ITIS |id=634389 |taxon=Scolecophidia |accessdate=14 August 2007}}</ref> are an [[infraorder]]<ref name="ITIS" /> of [[snakes]].<ref name="Vitt2014">{{Cite book|title=Herpetology: An Introductory Biology of Amphibians and Reptiles|last1=Vitt|first1=Laurie J.|last2=Caldwell|first2=Janalee P.|publisher=Academic Press|year=2014|isbn=978-0-12-386919-7|pages=597–603|edition=Fourth}}</ref> They range in length from {{convert|10|to|100|cm|in|sigfig=1|abbr=off}}. All are [[fossorial]] (adapted for burrowing).<ref>[http://palaeos.com/vertebrates/squamata/scolecophidia.html Scolecophidia] at [http://www.palaeos.com/ Palaeos]. Accessed 21 December 2013.</ref> Five families and 39 genera are recognized.<ref name="TRD">{{cite web|last1=Uetz|first1=Peter|title=The Reptile Database|url=http://www.reptile-database.reptarium.cz/advanced_search|website=The Reptile Database|accessdate=31 December 2017}}</ref> The Scolecophidia infraorder is almost likely [[Paraphyly|paraphyletic]]<ref>{{cite journal |last1=Miralles |first1=Aurélien |last2=Marin |first2=Julie |last3=Markus |first3=Damin |last4=Herrel |first4=Anthony |last5=Hedges |first5=Blair |last6=Vidal |first6=Nicolas |title=Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications |journal=Journal of Evolutionary Biology |date=2018 |volume=31 |issue=12 |pages=1782–1793 |doi=10.1111/jeb.13373|pmid=30193402 |s2cid=52174313 }}</ref>.
+The '''Scolecophidia''', commonly known as '''blind snakes''' or '''thread snakes''',<ref name="ITIS">{{ITIS |id=634389 |taxon=Scolecophidia |accessdate=14 August 2007}}</ref> are an [[infraorder]]<ref name="ITIS" /> of [[snakes]].<ref name="Vitt2014">{{Cite book|title=Herpetology: An Introductory Biology of Amphibians and Reptiles|last1=Vitt|first1=Laurie J.|author1-link=species:Laurie Joseph Vitt|last2=Caldwell|first2=Janalee P.|author2-link=species:Janalee Paige Caldwell|publisher=Academic Press|year=2014|isbn=978-0-12-386919-7|pages=597–603|edition=Fourth}}</ref> They range in length from {{convert|10|to|100|cm|in|sigfig=1|abbr=off}}. All are [[fossorial]] (adapted for burrowing).<ref>[http://palaeos.com/vertebrates/squamata/scolecophidia.html Scolecophidia] at [http://www.palaeos.com/ Palaeos]. Accessed 21 December 2013.</ref> Five [[Family (biology)|families]] and 39 [[genus|genera]] are recognized.<ref name="TRD">{{cite web|last1=Uetz|first1=Peter|author1-link=species:Peter Uetz|title=The Reptile Database|url=http://www.reptile-database.reptarium.cz/advanced_search|website=The Reptile Database|access-date=31 December 2017}}</ref> The Scolecophidia infraorder is most likely [[Paraphyly|paraphyletic]] (with the family [[Anomalepididae]] recovered with strong support as sister clade to the [[Alethinophidia|'typical snakes']]).<ref>{{cite journal |last1=Miralles |first1=Aurélien |author1-link=species:Aurélien Miralles |last2=Marin |first2=Julie |author2-link=species:Julie Marin |last3=Markus |first3=Damin |last4=Herrel |first4=Anthony |author4-link=species:Anthony Herrel |last5=Hedges |first5=Blair |author5-link=Stephen Blair Hedges |last6=Vidal |first6=Nicolas |author6-link=species:Nicolas Vidal |title=Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications |journal=Journal of Evolutionary Biology |date=2018 |volume=31 |issue=12 |pages=1782–1793 |doi=10.1111/jeb.13373|pmid=30193402 |s2cid=52174313 |doi-access=free }}</ref>
-== Systematics ==
+== Taxonomy==
-=== Etymology ===
 The infraorder name Scolecophidia derives from the two [[Ancient Greek]] words {{wikt-lang|grc|σκώληξ}} or σκώληκος ({{grc-transl|σκώληξ}}, genitive {{grc-transl|σκώληκος}}), meaning "earthworm", and {{wikt-lang|grc|ὄφις}} ({{grc-transl|ὄφις}}), meaning "snake".<ref>{{Cite book |title=Abrégé du dictionnaire grec français |last=Bailly |first=Anatole |date=1981-01-01 |publisher=Hachette |isbn=978-2010035289 |location=Paris |oclc=461974285 }}</ref><ref>{{Cite web |url=http://www.tabularium.be/bailly/ |title=Greek-french dictionary online |last=Bailly |first=Anatole |website=www.tabularium.be |access-date=January 7, 2019}}</ref> It refers to their shape and fossorial lifestyle.
@@ Line 31: / Line 30: @@
 |-
 |[[Gerrhopilidae]]
-|[[Nicolas Vidal|Vidal]], [[Addison H. Wynn|Wynn]], [[Steve Donnellan (scientist)|Donnellan]] & [[Stephen Blair Hedges|Hedges]], 2010<ref name="Vidal-2010">{{Cite journal|author=Vidal, Nicolas|year=2010|title=Blindsnake evolutionary tree reveals long history on Gondwana|journal=Biology Letters|volume=6|issue=4|pages=558–561, page 560|doi=10.1098/rsbl.2010.0220|display-authors=etal|pmid=20356885|pmc=2936224}}</ref>
+|[[species:Nicolas Vidal|Vidal]], [[species:Addison H. Wynn|Wynn]], [[Steve Donnellan (scientist)|Donnellan]] & [[Stephen Blair Hedges|Hedges]], 2010<ref name="Vidal-2010">{{Cite journal|author=Vidal, Nicolas|year=2010|title=Blindsnake evolutionary tree reveals long history on Gondwana|journal=Biology Letters|volume=6|issue=4|pages=558–561, page 560|doi=10.1098/rsbl.2010.0220|display-authors=etal|pmid=20356885|pmc=2936224}}</ref>
 |align="center"|2
-|blind snakes
+|Indo-Malayan blind snakes
 |style="width:40%"| [[India]], [[Southeast Asia]], [[Indonesia]], the [[Philippines]], and [[New Guinea]]<ref>{{cite book|author1=Vitt, Laurie J.|author2=Caldwell, Janalee P.|title=Herpetology: An Introductory Biology of Amphibians and Reptiles|publisher=Academic Press|year=2013|pages=600|url=https://books.google.com/books?id=Gay9N_ry79kC|isbn=9780123869203}}</ref>
 |-
@@ Line 51: / Line 50: @@
 |Vidal, [[Miguel Vences|Vences]], [[:fr:William Roy Branch|Branch]] & Hedges, 2010<ref name="Vidal-2010" />
 |align="center"|1
-|blind snakes
+|Malagasy blind snakes
 |style="width:40%"| [[Madagascar]]
 |-
 |}
-=== Chronology ===
+=== Evolution ===
 Despite only having fossils as early as the [[Cretaceous]], Scolecophidia itself likely originated in the [[Middle Jurassic]], with [[Anomalepididae]], [[Leptotyphlopidae]], and [[Typhlopoidea]] diverging from one another during the [[Late Jurassic]]. Within Typhlopoidea, [[Gerrhopilidae]] likely diverged from the Xenotyphlopidae-Typhlopidae [[clade]] during the [[Early Cretaceous]], and [[Xenotyphlopidae]] and [[Typhlopidae]] likely diverged from one another during the [[Late Cretaceous]].<ref name="Vidal-2010" />
+Scolecophidians are believed to have originated on [[Gondwana]], 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 [[Indian subcontinent|India]]/[[Madagascar]] land mass, during the [[Mesozoic]].<ref name="Vidal-2010"/>  Typhlopids, initially isolated on Madagascar, then dispersed to Africa and Eurasia. South American typhlopids appear to have evolved from African typhlopids that [[oceanic dispersal|rafted]] across the Atlantic about 60 million years ago; they, in turn, dispersed to the Caribbean about 33 million years ago.<ref name="Vidal-2010"/> Similarly, typhlopids appear to have reached Australia from Southeast Asia or Indonesia about 28 million years ago.<ref name="Vidal-2010"/> Meanwhile, the gerrhopilids, isolated on [[Insular India]], underwent a radiation throughout tropical Asia following the [[Paleogeography of the India–Asia collision system|collision of India with Asia]], while the xenotyphlopids remained isolated on Madagascar.<ref name=":1">{{Cite journal |last=Ali |first=Jason R. |last2=Hedges |first2=S. Blair |date=2023-05-04 |title=The colonisation of Madagascar by land‐bound vertebrates |url=https://onlinelibrary.wiley.com/doi/10.1111/brv.12966 |journal=Biological Reviews |language=en |doi=10.1111/brv.12966 |issn=1464-7931}}</ref>
+The Malagasy typhlopoids (''[[Madatyphlops]]'' in Typhlopidae and ''[[Xenotyphlops]]'' in Xenotyphlopidae) are among the only extant terrestrial vertebrates on Madagascar whose isolation occurred due to [[Allopatric speciation|vicariance]] 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.<ref name=":1" />
 === Fossil record ===
+[[File:Cobra-cega.jpg|thumb|Illustration of ''[[Boipeba tayasuensis|Boipeba]]'', the earliest known fossil blind snake]]
-The extinct fossil species ''[[Boipeba|Boipeba tayasuensis]]'' from the [[Late Cretaceous]] of [[Brazil]] was described in 2020, marking the earliest fossil record of Scolecophidia. It was a [[sister group]] to [[Typhlopoidea]] and was over 1 meter in length, making it much larger than most modern blindsnakes, with only ''[[Afrotyphlops schlegelii]]'' and ''[[Afrotyphlops mucruso]]'' rivaling it in size. Prior to this, the earliest scolecophidian fossils were only known from the [[Paleocene]] of [[Morocco]] and the [[Eocene]] of [[Europe]].<ref name=":0">{{Cite journal|author1=Thiago Schineider Fachini|author2=Silvio Onary|author3=Alessandro Palci|author4=Michael S.Y. Lee|author5=Mario Bronzati|author6=Annie Schmaltz Hsiou|year=2020|title=Cretaceous blindsnake from Brazil fills major gap in snake evolution|journal=iScience|volume=in press|pages=Article 101834|doi=10.1016/j.isci.2020.101834}}</ref>
+The extinct fossil species ''[[Boipeba tayasuensis]]'' from the [[Late Cretaceous]] of [[Brazil]] was described in 2020, marking the earliest fossil record of Scolecophidia. It was a [[sister group]] to [[Typhlopoidea]] and was over 1 meter in length, making it much larger than most modern blindsnakes, with only ''[[Afrotyphlops schlegelii]]'' and ''[[Afrotyphlops mucruso]]'' rivaling it in size. Prior to this, the earliest scolecophidian fossils were only known from the [[Paleocene]] of [[Morocco]] and the [[Eocene]] of [[Europe]].<ref name=":0">{{Cite journal|author1=Thiago Schineider Fachini|author2=Silvio Onary|author3=Alessandro Palci|author4=Michael S.Y. Lee|author4-link=species:Michael S.Y. Lee|author5=Mario Bronzati|author5-link=species:Mario Bronzati Filho|author6=Annie Schmaltz Hsiou|year=2020|title=Cretaceous blindsnake from Brazil fills major gap in snake evolution|journal=iScience|volume=23|issue=12|pages=Article 101834|doi=10.1016/j.isci.2020.101834|pmid=33305189|pmc=7718481|doi-access=free}}</ref>
+Possible Typhopid skin has been identified in [[Dominican amber]].<ref>{{Cite book|last1=Poinar|first1=George O.|url=https://books.google.com/books?id=MUJu9_zrPLQC|title=The Amber Forest: A Reconstruction of a Vanished World|last2=Poinar|first2=Roberta|date=1999|publisher=Princeton University Press|isbn=978-0-691-05728-6|language=en}}</ref>
 === Phylogeny ===
@@ Line 69: / Line 75: @@
     |1=[[Leptotyphlopidae]]
     |2={{clade
-      |1={{extinct}}''[[Boipeba]]''
+      |1={{extinct}}''[[Boipeba tayasuensis|Boipeba]]''
-      |label2=[[Typhlopoidea]]
+      |label2=[[Typhlopoidea (Superfamily)|Typhlopoidea]]
       |2={{clade
            |1=[[Gerrhopilidae]]
@@ Line 82: / Line 88: @@
 }}|label1='''Scolecophidia'''}}
-== Biogeographic history ==
+== Description ==
+The common name of Scolecophidia, blind snakes, is based on their shared characteristic of reduced eyes that are located under their head scales.<ref name="Simões2015">{{Cite journal|last=Simões|first=B. F. |display-authors=etal |date=2015|title=Visual system evolution and the nature of the ancestral snake|journal=Journal of Evolutionary Biology|volume=28|issue=7|pages=1309–1320|doi=10.1111/jeb.12663|pmid=26012745 |s2cid=24013194 |doi-access=free}}</ref> 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.<ref name="Vitt2014" /> 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.<ref name="Simões2015" /> 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.<ref>[https://www.sciencedaily.com/releases/2021/12/211209082611.htm 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.<ref name="Vitt2014" /> Aside from this, these snakes range in length from {{cvt|10|to|100|cm|0}}. Their typical body shapes include slender, cylindrical bodies and small, narrow heads.<ref name="Simões2015" /> All these families either lack or have a vestigial left lung and lack cranial infrared receptors.<ref name="Vitt2014" />
-Scolecophidians are believed to have originated on [[Gondwana]], with anomalepidids and leptotyphlopids evolving in west Gondwana (South America and Africa) and typhlopids, gerrhopilids, and xenotyphlopids on east Gondwana, initially on the combined India/Madagascar land mass, during the [[Mesozoic]].<ref name="Vidal-2010"/>  Typhlopids then dispersed to Africa and Eurasia. South American typhlopids appear to have evolved from African typhlopids that [[oceanic dispersal|rafted]] across the Atlantic about 60 million years ago; they, in turn, dispersed to the Caribbean about 33 million years ago.<ref name="Vidal-2010"/> Similarly, typhlopids appear to have reached Australia from Southeast Asia or Indonesia about 28 million years ago.<ref name="Vidal-2010"/>
-== Physical description ==
-The common name of Scolecophidia, blind snakes, is based on their shared characteristic of reduced eyes that are located under their head scales.<ref name="Simões2015">{{Cite journal|last=Simões|first=B. F. |display-authors=etal |date=2015|title=Visual system evolution and the nature of the ancestral snake|journal=Journal of Evolutionary Biology|volume=28|issue=7|pages=1309–1320|doi=10.1111/jeb.12663|pmid=26012745 |s2cid=24013194 }}</ref> 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.<ref name="Vitt2014" /> 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.<ref name="Simões2015" />  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.<ref name="Vitt2014" /> Aside from this, these snakes range in length from 10 to 100&nbsp;cm (3.9 to 39.4 in). Their typical body shapes include slender, cylindrical bodies and small, narrow heads.<ref name="Simões2015" /> All these families either lack or have a vestigial left lung and lack cranial infrared receptors.<ref name="Vitt2014" />
 == Behavior ==
-The main shared characteristic found across all Scolecophidia is a fossorial nature, either living underground or within logs and leaf litter.<ref name="Vitt2014" /> Aside from this, thus far the reproduction remains understudied with all Scolecophidia studied thus far being noted to be oviparous,<ref name="Vitt2014" /> with elongate eggs noted in both leptotyphlopids and typhlopids.<ref name="Webb2000">{{Cite journal|last1=Webb|first1=J. K.|last2=Shine|first2=R.|last3=Branch|first3=W. R.|last4=Harlow|first4=P. S.|date=2000|title=Life‐history strategies in basal snakes: reproduction and dietary habits of the African thread snake Leptotyphlops scutifrons (Serpentes: Leptotyphlopidae)|journal=Journal of Zoology|volume=250|issue=3|pages=321–327|doi=10.1017/s0952836900003058}}</ref> 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.<ref name="Webb2000" /> ''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.<ref name="Vitt2014" /> 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.<ref name="Vitt2014" /> 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.<ref name="Webb2000" />
+The main shared characteristic found across all Scolecophidia is a fossorial nature, either living underground or within logs and leaf litter.<ref name="Vitt2014" /> Aside from this, thus far the reproduction remains understudied with all Scolecophidia studied thus far being noted to be oviparous,<ref name="Vitt2014" /> with elongate eggs noted in both leptotyphlopids and typhlopids.<ref name="Webb2000">{{Cite journal|last1=Webb|first1=J.K.|author1-link=species:Jonathan K. Webb|last2=Shine|first2=R.|author2-link=Richard Shine|last3=Branch|first3=W.R.|author3-link=William Roy Branch|last4=Harlow|first4=P.S.|author4-link=species:Peter S. Harlow|date=2000|title=Life‐history strategies in basal snakes: reproduction and dietary habits of the African thread snake ''Leptotyphlops scutifrons'' (Serpentes: Leptotyphlopidae)|journal=Journal of Zoology|volume=250|issue=3|pages=321–327|doi=10.1017/s0952836900003058}}</ref> 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.<ref name="Webb2000" /> ''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.<ref name="Vitt2014" /> 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.<ref name="Vitt2014" /> 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.<ref name="Webb2000" />
-== See also ==
-*[[Alethinophidia]], all other snakes
-*[[List of snakes]], overview of all snake genera
 == References ==
@@ Line 102: / Line 100: @@
 {{Commons}}
+{{Squamata families}}
 {{Snake families}}
 {{Taxonbar|from=Q201636}}
@@ Line 107: / Line 106: @@
 [[Category:Scolecophidia| ]]
 [[Category:Taxa named by Edward Drinker Cope]]
-[[Category:Extant Late Cretaceous first appearances]]
+[[Category:Extant Coniacian first appearances]]

v t e Snake families
Kingdom: Animalia Phylum: Chordata Class: Reptilia Order: Squamata
Alethinophidia	Acrochordidae Aniliidae Anomochilidae Atractaspididae Boidae Bolyeriidae Colubridae Cyclocoridae Cylindrophiidae Elapidae Homalopsidae Lamprophiidae Loxocemidae Micrelapidae Prosymnidae Psammodynastidae Psammophiidae Pseudaspididae Pseudoxyrhophiidae Pythonidae Tropidophiidae Uropeltidae Viperidae Xenodermidae Xenopeltidae Xenophidiidae
Scolecophidia	Anomalepididae Gerrhopilidae Leptotyphlopidae Typhlopidae Xenotyphlopidae