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{{Short description|Subfamily of snakes}}
{{Short description|Subfamily of snakes}}
{{Automatic taxobox
{{Automatic taxobox
| fossil_range = {{fossil range|Upper Pleistocene|Recent}} <ref>{{cite web |title=Subfamily Dipsadinae Guenther 1858 (colubrid snake) |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=67685&is_real_user=1 |website=PBDB}}</ref>
| fossil_range = {{fossil range|Miocene|recent}}<ref>{{cite web |title=Xenodontinae |url=https://www.mindat.org/taxon-P53079.html|website=Mindat.org}}</ref>
| image = Sibon longifrenis (La Selva Biological Station).jpg
| image = Sibon longifrenis (La Selva Biological Station).jpg
| image_caption = ''[[Sibon longifrenis]]''
| image_caption = ''[[Sibon longifrenis]]''
| taxon = Dipsadinae
| taxon = Dipsadinae
| authority = [[Charles Lucien Bonaparte|Bonaparte]], 1838
| authority = [[Charles Lucien Bonaparte|Bonaparte]], 1838
| synonyms = Dipsadina <small>Bonaparte, 1838</small><ref name=Bonaparte/><br />
| synonyms =
Dipsadina <small>Bonaparte, 1838</small><ref name=Bonaparte/><br />
Dipsadidae <small>Bonaparte, 1838</small>
Dipsadidae <small>Bonaparte, 1838</small>
}}
}}


'''Dipsadinae''' is a large subfamily of [[Colubroidea|colubroid]] [[snakes]], sometimes referred to as a family ('''Dipsadidae''').<ref>{{cite journal|last1=Pyron|first1=R. A.|last2=Burbrink|first2=F.|last3=Wiens|first3=J. J.|title=A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes|journal=BMC Evolutionary Biology|date=2013|volume=13|pages=93|doi=10.1186/1471-2148-13-93|pmid=23627680|pmc=3682911}}</ref><ref name="Pyron2011">{{cite journal|last1=Pyron|first1=R. A.|last2=Burbrink|first2=F. T.|last3=Colli|first3=G. R.|last4=De Oca|first4=A. N. M.|last5=Vitt|first5=L. J.|last6=Kuczynski|first6=C. A.|last7=Wiens|first7=J. J.|title=The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees|journal=Molecular Phylogenetics and Evolution|date=2011|volume=58|issue=2|pages=329–342|doi=10.1016/j.ympev.2010.11.006|pmid=21074626|url=http://www.wienslab.com/Publications_files/Pyron_et_al_2011.pdf|quote=Our results support monophyly of Colubridae, containing the traditionally recognized subfamilies Calamariinae, Colubrinae, Natricinae, Pseudoxenodontinae, and Dipsadinae.}}</ref><ref name="Figueroa16">{{cite journal|last1=Figueroa|first1=A.|last2=McKelvy|first2=A. D.|last3=Grismer|first3=L. L.|last4=Bell|first4=C. D.|last5=Lailvaux|first5=S. P.|title=A species-level phylogeny of extant snakes with description of a new colubrid subfamily and genus|journal=PLOS ONE|date=2016|volume=11|issue=9|pages=e0161070|doi=10.1371/journal.pone.0161070|pmid=27603205|pmc=5014348|bibcode=2016PLoSO..1161070F |doi-access=free}}</ref><ref name="ZhengWiens2016">{{cite journal|last1=Zheng|first1=Y|last2=Wiens|first2=JJ|title=Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species|journal=Molecular Phylogenetics and Evolution|date=2016|volume=94|issue=Pt B|pages=537–547|url=http://www.wienslab.com/Publications_files/Zheng_Wiens_2015b_MPE.pdf|pmid=26475614|doi=10.1016/j.ympev.2015.10.009}}</ref><ref name=TRD/> They are found in most of the [[Americas]], including the [[West Indies]], and are most diverse in South America.<ref name="Grazziotin et al. 2012"/><ref name=VittCaldwell/> There are more than 700 species.<ref name="TRD">{{cite web|last1=Uetz|first1=Peter|title=Dipsadinae|url=http://reptile-database.reptarium.cz/advanced_search?taxon=dipsadinae&submit=Search|website=The Reptile Database|access-date=14 May 2018}}</ref>
'''Dipsadinae''' is a large [[subfamily]] of [[Colubroidea|colubroid]] [[snakes]], sometimes referred to as a [[Family (biology)|family]] ('''Dipsadidae''').<ref>{{cite journal|last1=Pyron|first1=R. A.|last2=Burbrink|first2=F.|last3=Wiens|first3=J. J.|title=A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes|journal=BMC Evolutionary Biology|date=2013|volume=13|pages=93|doi=10.1186/1471-2148-13-93|pmid=23627680|pmc=3682911 |doi-access=free }}</ref><ref name="Pyron2011">{{cite journal|last1=Pyron|first1=R. A.|last2=Burbrink|first2=F. T.|last3=Colli|first3=G. R.|last4=De Oca|first4=A. N. M.|last5=Vitt|first5=L. J.|last6=Kuczynski|first6=C. A.|last7=Wiens|first7=J. J.|title=The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees|journal=Molecular Phylogenetics and Evolution|date=2011|volume=58|issue=2|pages=329–342|doi=10.1016/j.ympev.2010.11.006|pmid=21074626|url=http://www.wienslab.com/Publications_files/Pyron_et_al_2011.pdf|quote=Our results support monophyly of Colubridae, containing the traditionally recognized subfamilies Calamariinae, Colubrinae, Natricinae, Pseudoxenodontinae, and Dipsadinae.}}</ref><ref name="Figueroa16">{{cite journal|last1=Figueroa|first1=A.|last2=McKelvy|first2=A. D.|last3=Grismer|first3=L. L.|last4=Bell|first4=C. D.|last5=Lailvaux|first5=S. P.|title=A species-level phylogeny of extant snakes with description of a new colubrid subfamily and genus|journal=PLOS ONE|date=2016|volume=11|issue=9|pages=e0161070|doi=10.1371/journal.pone.0161070|pmid=27603205|pmc=5014348|bibcode=2016PLoSO..1161070F |doi-access=free}}</ref><ref name="ZhengWiens2016">{{cite journal|last1=Zheng|first1=Y|last2=Wiens|first2=JJ|title=Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species|journal=Molecular Phylogenetics and Evolution|date=2016|volume=94|issue=Pt B|pages=537–547|url=http://www.wienslab.com/Publications_files/Zheng_Wiens_2015b_MPE.pdf|pmid=26475614|doi=10.1016/j.ympev.2015.10.009}}</ref><ref name=TRD/> [[Species]] of the subfamily Dipsadinae are found in most of the [[Americas]], including the [[West Indies]], and are most diverse in [[South America]].<ref name="Grazziotin et al. 2012"/><ref name=VittCaldwell/> There are more than 700 member species.<ref name="TRD">{{cite web|last1=Uetz|first1=Peter|title=Dipsadinae|url=http://reptile-database.reptarium.cz/advanced_search?taxon=dipsadinae&submit=Search|website=The Reptile Database|access-date=14 May 2018}}</ref>


Dipsadinae are an ecologically and morphologically diverse group of mostly small to moderate-sized snakes (typically less than {{convert|80|cm|abbr=on}} in total length). Some are arboreal, but others are aquatic or terrestrial and may even burrow. Most are [[oviparous]].<ref name=VittCaldwell/> Many eat frogs or lizards, and some consume mammals and birds. Several genera (e.g. ''[[Adelphicos]]'', ''[[Atractus]]'', ''[[Geophis]]'', ''[[Dipsas]]'', ''[[Ninia]]'', ''[[Sibon (snake)|Sibon]]'', ''[[Sibynomorphus]]'', ''[[Tropidodipsas]]'') are specialized feeders on gooey and slimy prey, such as frog eggs, earthworms, snails, and slugs.<ref name="Ray2012">{{cite journal|last1=Ray|first1=J. M.|last2=Montgomery|first2=C. E.|last3=Mahon|first3=H. K.|last4=Savitzky|first4=A. H.|last5=Lips|first5=K. R.|title=Goo-eaters: diets of the neotropical snakes Dipsas and Sibon in Central Panama|journal=Copeia|date=2012|volume=2012|issue=2|pages=197–202|doi=10.1643/CH-10-100|s2cid=86226277}}</ref><ref name="deOliveira08">{{cite journal|last1=de Oliveira|first1=L.|last2=Jared|first2=C.|last3=da Costa Prudente|first3=A. L.|last4=Zaher|first4=H.|last5=Antoniazzi|first5=M. M.|title=Oral glands in dipsadine "goo-eater" snakes: morphology and histochemistry of the infralabial glands in Atractus reticulatus, Dipsas indica, and Sibynomorphus mikanii|journal=Toxicon|date=2008|volume=51|issue=5|pages=898–913|doi=10.1016/j.toxicon.2007.12.021|pmid=18262581}}</ref><ref name="Zaher14">{{cite journal|last1=Zaher|first1=H.|last2=de Oliveira|first2=L.|last3=Grazziotin|first3=F. G.|last4=Campagner|first4=M.|last5=Jared|first5=C.|last6=Antoniazzi|first6=M. M.|last7=Prudente|first7=A. L.|title=Consuming viscous prey: a novel protein-secreting delivery system in neotropical snail-eating snakes|journal=BMC Evolutionary Biology|date=2014|volume=14|issue=1|pages=58|doi=10.1186/1471-2148-14-58|pmid=24661572|pmc=4021269}}</ref><ref>{{Cite web |url=http://faculty.weber.edu/rmeyers/lip-curling.pdf |title=Archived copy |access-date=2018-11-30 |archive-date=2018-12-01 |archive-url=https://web.archive.org/web/20181201005239/http://faculty.weber.edu/rmeyers/lip-curling.pdf |url-status=dead }}</ref><ref>{{Cite book | url=https://books.google.com/books?id=5XhvDwAAQBAJ |title = The Book of Snakes: A Life-Size Guide to Six Hundred Species from around the World|isbn = 9780226459394|last1 = O'Shea|first1 = Mark|date = 2018-10-22}}</ref> Almost all species are completely harmless to humans, although a few genera (e.g. ''[[Borikenophis]]'', ''[[Cubophis]]'', ''[[Heterodon]]'', ''[[Hydrodynastes]]'', ''[[Philodryas]]'') have inflicted painful bites with local, non-life-threatening symptoms.<ref name="Weinstein2011">{{cite book|last1=Weinstein|first1=S. A.|last2=Warrell|first2=D. A.|last3=White|first3=J.|last4=Keyler|first4=D. E.|title=Venomous bites from non-venomous snakes: A critical analysis of risk and management of "colubrid" snake bites|date=2011|publisher=Elsevier|location=London}}</ref>
Dipsadinae are an ecologically and morphologically diverse group of mostly small to moderate-sized snakes, typically less than {{convert|80|cm|abbr=on}} in total length. Some are arboreal, but others are aquatic or terrestrial and may even burrow. Most are [[oviparous]].<ref name=VittCaldwell/> Many eat frogs or lizards, and some consume mammals and birds. Several genera (e.g. ''[[Adelphicos]]'', ''[[Atractus]]'', ''[[Geophis]]'', ''[[Dipsas]]'', ''[[Ninia]]'', ''[[Sibon (snake)|Sibon]]'', ''[[Sibynomorphus]]'', ''[[Tropidodipsas]]'') are specialized feeders on gooey and slimy prey, such as frog eggs, earthworms, snails, and slugs.<ref name="Ray2012">{{cite journal|last1=Ray|first1=J. M.|last2=Montgomery|first2=C. E.|last3=Mahon|first3=H. K.|last4=Savitzky|first4=A. H.|last5=Lips|first5=K. R.|title=Goo-eaters: diets of the neotropical snakes Dipsas and Sibon in Central Panama|journal=Copeia|date=2012|volume=2012|issue=2|pages=197–202|doi=10.1643/CH-10-100|s2cid=86226277}}</ref><ref name="deOliveira08">{{cite journal|last1=de Oliveira|first1=L.|last2=Jared|first2=C.|last3=da Costa Prudente|first3=A. L.|last4=Zaher|first4=H.|last5=Antoniazzi|first5=M. M.|title=Oral glands in dipsadine "goo-eater" snakes: morphology and histochemistry of the infralabial glands in Atractus reticulatus, Dipsas indica, and Sibynomorphus mikanii|journal=Toxicon|date=2008|volume=51|issue=5|pages=898–913|doi=10.1016/j.toxicon.2007.12.021|pmid=18262581}}</ref><ref name="Zaher14">{{cite journal|last1=Zaher|first1=H.|last2=de Oliveira|first2=L.|last3=Grazziotin|first3=F. G.|last4=Campagner|first4=M.|last5=Jared|first5=C.|last6=Antoniazzi|first6=M. M.|last7=Prudente|first7=A. L.|title=Consuming viscous prey: a novel protein-secreting delivery system in neotropical snail-eating snakes|journal=BMC Evolutionary Biology|date=2014|volume=14|issue=1|pages=58|doi=10.1186/1471-2148-14-58|pmid=24661572|pmc=4021269 |doi-access=free }}</ref><ref>{{Cite web |url=http://faculty.weber.edu/rmeyers/lip-curling.pdf |title=Archived copy |access-date=2018-11-30 |archive-date=2018-12-01 |archive-url=https://web.archive.org/web/20181201005239/http://faculty.weber.edu/rmeyers/lip-curling.pdf |url-status=dead }}</ref><ref>{{Cite book | url=https://books.google.com/books?id=5XhvDwAAQBAJ |title = The Book of Snakes: A Life-Size Guide to Six Hundred Species from around the World|isbn = 9780226459394|last1 = O'Shea|first1 = Mark|date = 2018-10-22| publisher=University of Chicago Press }}</ref> Almost all species are completely harmless to humans, although a few genera (e.g. ''[[Borikenophis]]'', ''[[Cubophis]]'', ''[[Heterodon]]'', ''[[Hydrodynastes]]'', ''[[Philodryas]]'') have inflicted painful bites with local, non-life-threatening symptoms.<ref name="Weinstein2011">{{cite book|last1=Weinstein|first1=S. A.|last2=Warrell|first2=D. A.|last3=White|first3=J.|last4=Keyler|first4=D. E.|title=Venomous bites from non-venomous snakes: A critical analysis of risk and management of "colubrid" snake bites|date=2011|publisher=Elsevier|location=London}}</ref>

== Evolution ==
Molecular evidence indicates that Dipsadidae originated in Asia during the [[Ypresian|Early Eocene]], about 50 Ma. Xenodontinae and Dipsadinae diverged once the family had reached [[Central America]] in the Middle Eocene, with the latter dispersing to trans-Andean South America in multiple waves between 38 and 20 Ma, whereas the former dispersed to cis-Andean South America about 39 Ma and entered the West Indies some 33 Ma.<ref>{{Cite journal |last=Serrano |first=Filipe C. |last2=Pontes‐Nogueira |first2=Matheus |last3=Sawaya |first3=Ricardo J. |last4=Alencar |first4=Laura R. V. |last5=Nogueira |first5=Cristiano C. |last6=Grazziotin |first6=Felipe G. |date=3 January 2024 |title=There and back again: when and how the world's richest snake family (Dipsadidae) dispersed and speciated across the Neotropical region |url=https://onlinelibrary.wiley.com/doi/10.1111/jbi.14790 |journal=[[Journal of Biogeography]] |language=en |volume=51 |issue=5 |pages=878–893 |doi=10.1111/jbi.14790 |issn=0305-0270 |access-date=10 September 2024 |via=Wiley Online Library}}</ref>


==Synonymy==
==Synonymy==
Some authors refer to part or all of this group as [[Xenodontinae]], but if the two names are used synonymously, Dipsadinae is the correct name because it is older.<ref name="Pyron2011"/> When Xenodontinae is used non-synonymously, it normally refers to the larger and more derived South American-Caribbean subclade containing the genus ''[[Xenodon]]'' and its relatives, whereas Dipsadinae ''sensu stricto'' is restricted to the smaller and more basal Central American subclade containing the genus ''[[Dipsas]]'' and its relatives.<ref name="Grazziotin et al. 2012"/><ref name="Zaher2009">{{cite journal|last1=Zaher|first1=H.|last2=Grazziotin|first2=F. G.|last3=Cadle|first3=J. E.|last4=Murphy|first4=R. W.|last5=Moura-Leite|first5=J. C.|last6=Bonatto|first6=S. L.|title=Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American Xenodontines: A revised classification and descriptions of new taxa|journal=Papéis Avulsos de Zoologia (São Paulo)|date=2009|volume=49|issue=11|pages=115–153|doi=10.1590/S0031-10492009001100001|doi-access=free}}</ref> Also, a third North American group (sometimes called "Carphophiinae") contains nine species in five genera at the base of the Dipsadinae (the "North American relicts" thought to have descended from the ancestors of dipsadines as they crossed from Asia to South America by way of North America; genera ''[[Heterodon]]'', ''[[Farancia]]'', ''[[Diadophis]]'', ''[[Carphophis]]'', and ''[[Contia]]'').<ref name="Pinou2004">{{cite journal|last1=Pinou|first1=T.|last2=Vicario|first2=S.|last3=Marschner|first3=M.|last4=Caccone|first4=A.|title=Relict snakes of North America and their relationships within Caenophidia, using likelihood-based Bayesian methods on mitochondrial sequences|journal=Molecular Phylogenetics and Evolution|date=2004|volume=32|issue=2|pages=563–574|doi=10.1016/j.ympev.2004.02.005|pmid=15223038|citeseerx=10.1.1.542.4840}}</ref>
Some authors refer to part or all of this group as [[Xenodontinae]], but if the two names are used synonymously, Dipsadinae is the correct name because it is older.<ref name="Pyron2011"/> When Xenodontinae is used non-synonymously, it normally refers to the larger and more derived South American-Caribbean subclade containing the genus ''[[Xenodon]]'' and its relatives, whereas Dipsadinae ''sensu stricto'' is restricted to the smaller and more basal Central American subclade containing the genus ''[[Dipsas]]'' and its relatives.<ref name="Grazziotin et al. 2012"/><ref name="Zaher2009">{{cite journal|last1=Zaher|first1=H.|last2=Grazziotin|first2=F. G.|last3=Cadle|first3=J. E.|last4=Murphy|first4=R. W.|last5=Moura-Leite|first5=J. C.|last6=Bonatto|first6=S. L.|title=Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American Xenodontines: A revised classification and descriptions of new taxa|journal=Papéis Avulsos de Zoologia|date=2009|volume=49|issue=11|pages=115–153|doi=10.1590/S0031-10492009001100001|doi-access=free|hdl=11449/71032|hdl-access=free}}</ref> Also, a third North American group (sometimes called "Carphophiinae") contains nine species in five genera at the base of the Dipsadinae (the "North American relicts" thought to have descended from the ancestors of dipsadines as they crossed from Asia to South America by way of North America; genera ''[[Heterodon]]'', ''[[Farancia]]'', ''[[Diadophis]]'', ''[[Carphophis]]'', and ''[[Contia]]'').<ref name="Pinou2004">{{cite journal|last1=Pinou|first1=T.|last2=Vicario|first2=S.|last3=Marschner|first3=M.|last4=Caccone|first4=A.|title=Relict snakes of North America and their relationships within Caenophidia, using likelihood-based Bayesian methods on mitochondrial sequences|journal=Molecular Phylogenetics and Evolution|date=2004|volume=32|issue=2|pages=563–574|doi=10.1016/j.ympev.2004.02.005|pmid=15223038|citeseerx=10.1.1.542.4840}}</ref>


== Genera ==
== Genera ==


Within the Dipsadinae, the three major groups/clades or subfamilies are the Central American group ("Dipsadinae" ''sensu stricto''), the South American + Caribbean group ("[[Xenodontinae]]"), and a small North American group (sometimes called the "Carphophiinae" or, incorrectly, "Heterodontinae").{{efn|"[[Bullhead shark|Heterodontinae]]" is a subfamily of sharks; this problem is discussed in Appendix I of Grazziotin, F. G., H. Zaher, R. W. Murphy, G. Scrocchi, M. A. Benavides, Y.-P. Zhang, and S. L. Bonattoh (2012):<ref>{{cite journal|doi=10.1111/j.1096-0031.2012.00393.x|title=Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): A reappraisal|journal=Cladistics|volume=28|issue=5|pages=437–459|year=2012|last1=Grazziotin|first1=Felipe G|last2=Zaher|first2=Hussam|last3=Murphy|first3=Robert W|last4=Scrocchi|first4=Gustavo|last5=Benavides|first5=Marco A|last6=Zhang|first6=Ya-Ping|last7=Bonatto|first7=Sandro L|pmid=34836446 |s2cid=84934386}}</ref> ''"The shark family Heterodontidae (based on the genus Heterodontus Blainville, 1816) dates from Gray (1851: 65), but its use as the snake family Heterodontidae (based on the genus Heterodon Latreille, 1801) dates from Bonaparte (1845) and it has not been used in the literature since. Thus both the genus and family names for snakes have priority over the sharks. However, the resurrection of the family name Heterodontidae for snakes (subfamily Heterodontinae in Vidal et al., 2007) causes unnecessary confusion owing to the long-standing use of the name for sharks (e.g. Compagno, 2002; Baldwin, 2005). Consequently, Rossman and Wilson (1965) and Zaher et al. (2009) argued that the family name should be applied only to sharks in the interest of maintaining nomenclatorial stability, a position that contrasts strongly with that of Vidal et al. (2007, 2010). According to Art. 52.2 of the Code, when two names ‘‘are homonyms, only the senior, as determined by the Principle of Priority, may be used as a valid name’’. We believe that if this clade of snakes continuously appears in phylogenetic studies, then it is desirable to petition the International Commission on Zoological Nomenclature to set aside use of the family name for the snakes in favor of the sharks in the interest of nomenclatorial stability. An alternative nomenclature would be to change the spelling of the shark family to Heterodontusidae. In any case, we suggest the North American relictual Xenodontinae (sensu Pinou, 1993; Pinou et al., 2004) should not be referred to as the subfamily Heterodontinae until a well defined nomenclatural resolution is obtained."'' See also Rossman, D. A. and L. D. Wilson (1965).<ref>{{cite book|author=D. A. Rossman|title=Comments on the Revival of the Colubrid Snake Subfamily Heterodontinae|url=https://books.google.com/books?id=J5T4jgEACAAJ|year=1965}}</ref>}}<ref name="Pinou2004"/> In addition, a number of snake genera are likely to be dipsadines based on their morphology and geographic range, but because of the absence of genetic data and information about their closest relatives, they are considered genera'' [[incertae sedis]]'' and are not currently placed in a subgroup of the Dipsadinae.
Within the Dipsadinae, the three major groups/clades or subfamilies are the Central American group ("Dipsadinae" ''sensu stricto''), the South American + Caribbean group ("[[Xenodontinae]]"), and a small North American group (sometimes called the "Carphophiinae" or, incorrectly, "Heterodontinae").{{efn|"[[Bullhead shark|Heterodontinae]]" is a subfamily of sharks; this problem is discussed in Appendix I of Grazziotin, F. G., H. Zaher, R. W. Murphy, G. Scrocchi, M. A. Benavides, Y.-P. Zhang, and S. L. Bonattoh (2012):<ref>{{cite journal|doi=10.1111/j.1096-0031.2012.00393.x|title=Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): A reappraisal|journal=Cladistics|volume=28|issue=5|pages=437–459|year=2012|last1=Grazziotin|first1=Felipe G|last2=Zaher|first2=Hussam|last3=Murphy|first3=Robert W|last4=Scrocchi|first4=Gustavo|last5=Benavides|first5=Marco A|last6=Zhang|first6=Ya-Ping|last7=Bonatto|first7=Sandro L|pmid=34836446 |s2cid=84934386|doi-access=free}}</ref> ''"The shark family Heterodontidae (based on the genus Heterodontus Blainville, 1816) dates from Gray (1851: 65), but its use as the snake family Heterodontidae (based on the genus Heterodon Latreille, 1801) dates from Bonaparte (1845) and it has not been used in the literature since. Thus both the genus and family names for snakes have priority over the sharks. However, the resurrection of the family name Heterodontidae for snakes (subfamily Heterodontinae in Vidal et al., 2007) causes unnecessary confusion owing to the long-standing use of the name for sharks (e.g. Compagno, 2002; Baldwin, 2005). Consequently, Rossman and Wilson (1965) and Zaher et al. (2009) argued that the family name should be applied only to sharks in the interest of maintaining nomenclatorial stability, a position that contrasts strongly with that of Vidal et al. (2007, 2010). According to Art. 52.2 of the Code, when two names ‘‘are homonyms, only the senior, as determined by the Principle of Priority, may be used as a valid name’’. We believe that if this clade of snakes continuously appears in phylogenetic studies, then it is desirable to petition the International Commission on Zoological Nomenclature to set aside use of the family name for the snakes in favor of the sharks in the interest of nomenclatorial stability. An alternative nomenclature would be to change the spelling of the shark family to Heterodontusidae. In any case, we suggest the North American relictual Xenodontinae (sensu Pinou, 1993; Pinou et al., 2004) should not be referred to as the subfamily Heterodontinae until a well defined nomenclatural resolution is obtained."'' See also Rossman, D. A. and L. D. Wilson (1965).<ref>{{cite book|author=D. A. Rossman|title=Comments on the Revival of the Colubrid Snake Subfamily Heterodontinae|url=https://books.google.com/books?id=J5T4jgEACAAJ|year=1965}}</ref>}}<ref name="Pinou2004"/> In addition, a number of snake genera are likely to be dipsadines based on their morphology and geographic range, but because of the absence of genetic data and information about their closest relatives, they are considered genera'' [[incertae sedis]]'' and are not currently placed in a subgroup of the Dipsadinae.


=== Central American clade ("Dipsadinae" ''sensu stricto'') ===
=== Central American clade ("Dipsadinae" ''sensu stricto'') ===
Line 39: Line 41:
* ''[[Ninia]]'' Baird & Girard, 1853
* ''[[Ninia]]'' Baird & Girard, 1853
* ''[[Plesiodipsas]]'' Harvey, Fuenmayor, Portilla & Rueda-Almonacid, 2008
* ''[[Plesiodipsas]]'' Harvey, Fuenmayor, Portilla & Rueda-Almonacid, 2008
* ''[[Pliocercus]]''
* ''[[Pliocercus]]'' Cope, 1860
* ''[[Pseudoleptodeira]]'' Taylor, 1938
* ''[[Pseudoleptodeira]]'' Taylor, 1938
* ''[[Rhadinaea]]'' Cope, 1863
* ''[[Rhadinaea]]'' Cope, 1863
Line 67: Line 69:
** ''[[Apographon]]'' Trevine, Grazziotin, Giraudo, Sallesbery‐Pinchera, Vianna, & Zaher, 2022
** ''[[Apographon]]'' Trevine, Grazziotin, Giraudo, Sallesbery‐Pinchera, Vianna, & Zaher, 2022
** ''[[Calamodontophis]]'' Amaral, 1963
** ''[[Calamodontophis]]'' Amaral, 1963
** ''[[Dryophylax]]'' Wagler, 1830
** ''[[Galvarinus]]'' Trevine et al., 2022
** ''[[Gomesophis]]'' Hoge & Mertens, 1959
** ''[[Gomesophis]]'' Hoge & Mertens, 1959
** ''[[Pseudotomodon]]'' Koslowski, 1896
** ''[[Mesotes]]'' Trevine et al., 2022
** ''[[Ptychophis]]'' Gomes, 1915
** ''[[Ptychophis]]'' Gomes, 1915
** ''[[Tachymenis]]'' Wiegmann, 1835
** ''[[Tachymenis]]'' Wiegmann, 1835
** ''[[Tachymenoides]]'' Trevine et al., 2022
** ''[[Thamnodynastes]]'' Wagler, 1830
** ''[[Thamnodynastes]]'' Wagler, 1830
** ''[[Tomodon]]'' A.M.C. Duméril, 1853
** ''[[Tomodon]]'' A.M.C. Duméril, 1853
** ''[[Zonateres]]'' Trevine, V. C., Grazziotin, F. G., Giraudo, A., Sallesbery‐Pinchera, N., Vianna, J. A., & Zaher, H. 2022
* '''Tribe [[Echinantherini]]''' Zaher, Grazziotin, Cadle, Murphy, de Moura-Leite & Bonatto, 2009
* '''Tribe [[Echinantherini]]''' Zaher, Grazziotin, Cadle, Murphy, de Moura-Leite & Bonatto, 2009
** ''[[Adelphostigma]]'' Abegg, Santos-Jr, Costa, Battilana, Graboski, Vianna, Azevedo, Fagundes, Castille, Prado, Bonatto, Zaher, & Grazziotin, 2008
** ''[[Adelphostigma]]'' Abegg, Santos-Jr, Costa, Battilana, Graboski, Vianna, Azevedo, Fagundes, Castille, Prado, Bonatto, Zaher, & Grazziotin, 2008
Line 85: Line 91:
* '''Tribe [[Pseudoboini]]''' Bailey, 1967
* '''Tribe [[Pseudoboini]]''' Bailey, 1967
** ''[[Boiruna]]'' Zaher, 1996
** ''[[Boiruna]]'' Zaher, 1996
** ''[[Clelia (snake)|Clelia]]'' Fitzinger, 1826
** ''[[Clelia (snake genus)|Clelia]]'' Fitzinger, 1826
** ''[[Drepanoides]]'' Dunn, 1928
** ''[[Drepanoides]]'' Dunn, 1928
** ''[[Mussurana (genus)|Mussurana]]'' Zaher, Grazziotin, Cadle, Murphy, de Moura-Leite & Bonatto, 2009
** ''[[Mussurana (genus)|Mussurana]]'' Zaher, Grazziotin, Cadle, Murphy, de Moura-Leite & Bonatto, 2009
Line 128: Line 134:
** ''[[Magliophis]]'' Zaher, Grazziotin, Cadle, Murphy, de Moura-Leite & Bonatto, 2009
** ''[[Magliophis]]'' Zaher, Grazziotin, Cadle, Murphy, de Moura-Leite & Bonatto, 2009
** ''[[Uromacer]]'' A.M.C. Duméril, Bibron & A.H.A. Duméril, 1854
** ''[[Uromacer]]'' A.M.C. Duméril, Bibron & A.H.A. Duméril, 1854
* '''Tribe ''incertae sedis'''''
** ''[[Paikwaophis]]'' Kok & Means, 2023
** ''[[Xenopholis]]'' Peters, 1869

}}
}}


Line 158: Line 168:
* ''[[Synophis]]'' Peracca 1896
* ''[[Synophis]]'' Peracca 1896
* ''[[Tantalophis]]'' Duellman, 1958
* ''[[Tantalophis]]'' Duellman, 1958
* ''[[Thermophis]]'' Malnate, 1953 (the only Asian genus, likely at the base of the entire clade or at the base of the Dipdadinae + [[Natricinae]] clade)<ref>{{Cite journal | author1 = He M | author2 =Feng JC |author3 = Liu SY | author4 = Guo P | author5 = Zhao EM | author-link5 = Er-mi Zhao | title = The phylogenetic position of ''Thermophis'' (Serpentes: Colubridae), an endemic snake from the Qinghai‐Xizang Plateau, China | doi = 10.1080/00222930802389825 | journal = Journal of Natural History | volume = 43 | issue = 7–8 | pages = 479–488| year = 2009 | s2cid =84653966 | url = http://210.75.237.14/bitstream/351003/19125/1/2009e0050h.pdf}}</ref><ref name=Huang>{{Cite journal | author1 = Huang S |author2 = Liu SY |author3 = Guo P |author4 = Zhang YP |author5 = Zhao EM | title = What are the closest relatives of the hot-spring snakes (Colubridae, ''Thermophis''), the relict species endemic to the Tibetan Plateau? | doi = 10.1016/j.ympev.2009.02.013 | journal = Molecular Phylogenetics and Evolution | volume = 51 | issue = 3 | pages = 438–446 | year = 2009 | pmid = 19249375| url = http://210.75.237.14/bitstream/351003/19127/1/2009e0111h.pdf}}</ref>}}
* ''[[Thermophis]]'' Malnate, 1953 (the only Asian genus, likely at the base of the entire clade or at the base of the Dipsadinae + [[Natricinae]] clade)<ref>{{Cite journal | author1 = He M | author2 = Feng JC | author3 = Liu SY | author4 = Guo P | author5 = Zhao EM | author-link5 = Er-mi Zhao | title = The phylogenetic position of ''Thermophis'' (Serpentes: Colubridae), an endemic snake from the Qinghai‐Xizang Plateau, China | doi = 10.1080/00222930802389825 | journal = Journal of Natural History | volume = 43 | issue = 7–8 | pages = 479–488 | year = 2009 | s2cid = 84653966 | url = http://210.75.237.14/bitstream/351003/19125/1/2009e0050h.pdf | access-date = 2018-05-14 | archive-date = 2015-12-22 | archive-url = https://web.archive.org/web/20151222082320/http://210.75.237.14/bitstream/351003/19125/1/2009e0050h.pdf | url-status = dead }}</ref><ref name=Huang>{{Cite journal | author1 = Huang S | author2 = Liu SY | author3 = Guo P | author4 = Zhang YP | author5 = Zhao EM | title = What are the closest relatives of the hot-spring snakes (Colubridae, ''Thermophis''), the relict species endemic to the Tibetan Plateau? | doi = 10.1016/j.ympev.2009.02.013 | journal = Molecular Phylogenetics and Evolution | volume = 51 | issue = 3 | pages = 438–446 | year = 2009 | pmid = 19249375 | url = http://210.75.237.14/bitstream/351003/19127/1/2009e0111h.pdf | access-date = 2018-05-14 | archive-date = 2013-10-03 | archive-url = https://web.archive.org/web/20131003165734/http://210.75.237.14/bitstream/351003/19127/1/2009e0111h.pdf | url-status = dead }}</ref>}}


==Notes==
==Notes==
Line 167: Line 177:
<ref name=Bonaparte>{{cite journal |last1=Bonaparte |first1=Charles Lucien |year=1838 |title=Synopsis vertebratorum systematis. Amphibiorum Tabula Analytica |journal=Nuovi Annali delle Scienze Naturali |volume=1 |pages=391–397 |url=https://archive.org/details/nuoviannalidelle01nuov }}</ref>
<ref name=Bonaparte>{{cite journal |last1=Bonaparte |first1=Charles Lucien |year=1838 |title=Synopsis vertebratorum systematis. Amphibiorum Tabula Analytica |journal=Nuovi Annali delle Scienze Naturali |volume=1 |pages=391–397 |url=https://archive.org/details/nuoviannalidelle01nuov }}</ref>


<ref name="Grazziotin et al. 2012">{{cite journal|last1=Grazziotin|first1=Felipe G.|last2=Zaher|first2=Hussam|last3=Murphy|first3=Robert W.|last4=Scrocchi|first4=Gustavo|last5=Benavides|first5=Marco A.|last6=Zhang|first6=Ya-Ping|last7=Bonatto|first7=Sandro L. |year=2012 |title=Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): a reappraisal|journal=Cladistics |volume=28 |issue=5 |pages=437–459 |doi=10.1111/j.1096-0031.2012.00393.x|pmid=34836446 |s2cid=84934386}}</ref>
<ref name="Grazziotin et al. 2012">{{cite journal|last1=Grazziotin|first1=Felipe G.|last2=Zaher|first2=Hussam|last3=Murphy|first3=Robert W.|last4=Scrocchi|first4=Gustavo|last5=Benavides|first5=Marco A.|last6=Zhang|first6=Ya-Ping|last7=Bonatto|first7=Sandro L. |year=2012 |title=Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): a reappraisal|journal=Cladistics |volume=28 |issue=5 |pages=437–459 |doi=10.1111/j.1096-0031.2012.00393.x|pmid=34836446 |s2cid=84934386|doi-access=free}}</ref>


<ref name=VittCaldwell>{{cite book|title=Herpetology: An Introductory Biology of Amphibians and Reptiles |edition=4th |first1=Laurie J. |last1=Vitt |first2=Janalee P. |last2=Caldwell |publisher=Academic Press |year=2014 |pages=622–626}}</ref>
<ref name=VittCaldwell>{{cite book|title=Herpetology: An Introductory Biology of Amphibians and Reptiles |edition=4th |first1=Laurie J. |last1=Vitt |first2=Janalee P. |last2=Caldwell |publisher=Academic Press |year=2014 |pages=622–626}}</ref>
Line 175: Line 185:
{{Taxonbar|from=Q143402}}
{{Taxonbar|from=Q143402}}


[[Category:Dipsadidae]]
[[Category:Dipsadinae]]
[[Category:Reptile families]]
[[Category:Snake families]]
[[Category:Taxa named by Charles Lucien Bonaparte]]
[[Category:Taxa named by Charles Lucien Bonaparte]]

Latest revision as of 19:52, 10 September 2024

Dipsadinae
Temporal range: Miocene–recent [1]
Sibon longifrenis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Serpentes
Family: Colubridae
Subfamily: Dipsadinae
Bonaparte, 1838
Synonyms

Dipsadina Bonaparte, 1838[2]
Dipsadidae Bonaparte, 1838

Dipsadinae is a large subfamily of colubroid snakes, sometimes referred to as a family (Dipsadidae).[3][4][5][6][7] Species of the subfamily Dipsadinae are found in most of the Americas, including the West Indies, and are most diverse in South America.[8][9] There are more than 700 member species.[7]

Dipsadinae are an ecologically and morphologically diverse group of mostly small to moderate-sized snakes, typically less than 80 cm (31 in) in total length. Some are arboreal, but others are aquatic or terrestrial and may even burrow. Most are oviparous.[9] Many eat frogs or lizards, and some consume mammals and birds. Several genera (e.g. Adelphicos, Atractus, Geophis, Dipsas, Ninia, Sibon, Sibynomorphus, Tropidodipsas) are specialized feeders on gooey and slimy prey, such as frog eggs, earthworms, snails, and slugs.[10][11][12][13][14] Almost all species are completely harmless to humans, although a few genera (e.g. Borikenophis, Cubophis, Heterodon, Hydrodynastes, Philodryas) have inflicted painful bites with local, non-life-threatening symptoms.[15]

Evolution

[edit]

Molecular evidence indicates that Dipsadidae originated in Asia during the Early Eocene, about 50 Ma. Xenodontinae and Dipsadinae diverged once the family had reached Central America in the Middle Eocene, with the latter dispersing to trans-Andean South America in multiple waves between 38 and 20 Ma, whereas the former dispersed to cis-Andean South America about 39 Ma and entered the West Indies some 33 Ma.[16]

Synonymy

[edit]

Some authors refer to part or all of this group as Xenodontinae, but if the two names are used synonymously, Dipsadinae is the correct name because it is older.[4] When Xenodontinae is used non-synonymously, it normally refers to the larger and more derived South American-Caribbean subclade containing the genus Xenodon and its relatives, whereas Dipsadinae sensu stricto is restricted to the smaller and more basal Central American subclade containing the genus Dipsas and its relatives.[8][17] Also, a third North American group (sometimes called "Carphophiinae") contains nine species in five genera at the base of the Dipsadinae (the "North American relicts" thought to have descended from the ancestors of dipsadines as they crossed from Asia to South America by way of North America; genera Heterodon, Farancia, Diadophis, Carphophis, and Contia).[18]

Genera

[edit]

Within the Dipsadinae, the three major groups/clades or subfamilies are the Central American group ("Dipsadinae" sensu stricto), the South American + Caribbean group ("Xenodontinae"), and a small North American group (sometimes called the "Carphophiinae" or, incorrectly, "Heterodontinae").[a][18] In addition, a number of snake genera are likely to be dipsadines based on their morphology and geographic range, but because of the absence of genetic data and information about their closest relatives, they are considered genera incertae sedis and are not currently placed in a subgroup of the Dipsadinae.

Central American clade ("Dipsadinae" sensu stricto)

[edit]

South American + Caribbean clade ("Xenodontinae")

[edit]

North American clade ("Carphophiinae")

[edit]

Genera incertae sedis

[edit]

Notes

[edit]
  1. ^ "Heterodontinae" is a subfamily of sharks; this problem is discussed in Appendix I of Grazziotin, F. G., H. Zaher, R. W. Murphy, G. Scrocchi, M. A. Benavides, Y.-P. Zhang, and S. L. Bonattoh (2012):[19] "The shark family Heterodontidae (based on the genus Heterodontus Blainville, 1816) dates from Gray (1851: 65), but its use as the snake family Heterodontidae (based on the genus Heterodon Latreille, 1801) dates from Bonaparte (1845) and it has not been used in the literature since. Thus both the genus and family names for snakes have priority over the sharks. However, the resurrection of the family name Heterodontidae for snakes (subfamily Heterodontinae in Vidal et al., 2007) causes unnecessary confusion owing to the long-standing use of the name for sharks (e.g. Compagno, 2002; Baldwin, 2005). Consequently, Rossman and Wilson (1965) and Zaher et al. (2009) argued that the family name should be applied only to sharks in the interest of maintaining nomenclatorial stability, a position that contrasts strongly with that of Vidal et al. (2007, 2010). According to Art. 52.2 of the Code, when two names ‘‘are homonyms, only the senior, as determined by the Principle of Priority, may be used as a valid name’’. We believe that if this clade of snakes continuously appears in phylogenetic studies, then it is desirable to petition the International Commission on Zoological Nomenclature to set aside use of the family name for the snakes in favor of the sharks in the interest of nomenclatorial stability. An alternative nomenclature would be to change the spelling of the shark family to Heterodontusidae. In any case, we suggest the North American relictual Xenodontinae (sensu Pinou, 1993; Pinou et al., 2004) should not be referred to as the subfamily Heterodontinae until a well defined nomenclatural resolution is obtained." See also Rossman, D. A. and L. D. Wilson (1965).[20]

References

[edit]
  1. ^ "Xenodontinae". Mindat.org.
  2. ^ Bonaparte, Charles Lucien (1838). "Synopsis vertebratorum systematis. Amphibiorum Tabula Analytica". Nuovi Annali delle Scienze Naturali. 1: 391–397.
  3. ^ Pyron, R. A.; Burbrink, F.; Wiens, J. J. (2013). "A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes". BMC Evolutionary Biology. 13: 93. doi:10.1186/1471-2148-13-93. PMC 3682911. PMID 23627680.
  4. ^ a b Pyron, R. A.; Burbrink, F. T.; Colli, G. R.; De Oca, A. N. M.; Vitt, L. J.; Kuczynski, C. A.; Wiens, J. J. (2011). "The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees" (PDF). Molecular Phylogenetics and Evolution. 58 (2): 329–342. doi:10.1016/j.ympev.2010.11.006. PMID 21074626. Our results support monophyly of Colubridae, containing the traditionally recognized subfamilies Calamariinae, Colubrinae, Natricinae, Pseudoxenodontinae, and Dipsadinae.
  5. ^ Figueroa, A.; McKelvy, A. D.; Grismer, L. L.; Bell, C. D.; Lailvaux, S. P. (2016). "A species-level phylogeny of extant snakes with description of a new colubrid subfamily and genus". PLOS ONE. 11 (9): e0161070. Bibcode:2016PLoSO..1161070F. doi:10.1371/journal.pone.0161070. PMC 5014348. PMID 27603205.
  6. ^ Zheng, Y; Wiens, JJ (2016). "Combining phylogenomic and supermatrix approaches, and a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52 genes and 4162 species" (PDF). Molecular Phylogenetics and Evolution. 94 (Pt B): 537–547. doi:10.1016/j.ympev.2015.10.009. PMID 26475614.
  7. ^ a b Uetz, Peter. "Dipsadinae". The Reptile Database. Retrieved 14 May 2018.
  8. ^ a b Grazziotin, Felipe G.; Zaher, Hussam; Murphy, Robert W.; Scrocchi, Gustavo; Benavides, Marco A.; Zhang, Ya-Ping; Bonatto, Sandro L. (2012). "Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): a reappraisal". Cladistics. 28 (5): 437–459. doi:10.1111/j.1096-0031.2012.00393.x. PMID 34836446. S2CID 84934386.
  9. ^ a b Vitt, Laurie J.; Caldwell, Janalee P. (2014). Herpetology: An Introductory Biology of Amphibians and Reptiles (4th ed.). Academic Press. pp. 622–626.
  10. ^ Ray, J. M.; Montgomery, C. E.; Mahon, H. K.; Savitzky, A. H.; Lips, K. R. (2012). "Goo-eaters: diets of the neotropical snakes Dipsas and Sibon in Central Panama". Copeia. 2012 (2): 197–202. doi:10.1643/CH-10-100. S2CID 86226277.
  11. ^ de Oliveira, L.; Jared, C.; da Costa Prudente, A. L.; Zaher, H.; Antoniazzi, M. M. (2008). "Oral glands in dipsadine "goo-eater" snakes: morphology and histochemistry of the infralabial glands in Atractus reticulatus, Dipsas indica, and Sibynomorphus mikanii". Toxicon. 51 (5): 898–913. doi:10.1016/j.toxicon.2007.12.021. PMID 18262581.
  12. ^ Zaher, H.; de Oliveira, L.; Grazziotin, F. G.; Campagner, M.; Jared, C.; Antoniazzi, M. M.; Prudente, A. L. (2014). "Consuming viscous prey: a novel protein-secreting delivery system in neotropical snail-eating snakes". BMC Evolutionary Biology. 14 (1): 58. doi:10.1186/1471-2148-14-58. PMC 4021269. PMID 24661572.
  13. ^ "Archived copy" (PDF). Archived from the original (PDF) on 2018-12-01. Retrieved 2018-11-30.{{cite web}}: CS1 maint: archived copy as title (link)
  14. ^ O'Shea, Mark (2018-10-22). The Book of Snakes: A Life-Size Guide to Six Hundred Species from around the World. University of Chicago Press. ISBN 9780226459394.
  15. ^ Weinstein, S. A.; Warrell, D. A.; White, J.; Keyler, D. E. (2011). Venomous bites from non-venomous snakes: A critical analysis of risk and management of "colubrid" snake bites. London: Elsevier.
  16. ^ Serrano, Filipe C.; Pontes‐Nogueira, Matheus; Sawaya, Ricardo J.; Alencar, Laura R. V.; Nogueira, Cristiano C.; Grazziotin, Felipe G. (3 January 2024). "There and back again: when and how the world's richest snake family (Dipsadidae) dispersed and speciated across the Neotropical region". Journal of Biogeography. 51 (5): 878–893. doi:10.1111/jbi.14790. ISSN 0305-0270. Retrieved 10 September 2024 – via Wiley Online Library.
  17. ^ Zaher, H.; Grazziotin, F. G.; Cadle, J. E.; Murphy, R. W.; Moura-Leite, J. C.; Bonatto, S. L. (2009). "Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American Xenodontines: A revised classification and descriptions of new taxa". Papéis Avulsos de Zoologia. 49 (11): 115–153. doi:10.1590/S0031-10492009001100001. hdl:11449/71032.
  18. ^ a b Pinou, T.; Vicario, S.; Marschner, M.; Caccone, A. (2004). "Relict snakes of North America and their relationships within Caenophidia, using likelihood-based Bayesian methods on mitochondrial sequences". Molecular Phylogenetics and Evolution. 32 (2): 563–574. CiteSeerX 10.1.1.542.4840. doi:10.1016/j.ympev.2004.02.005. PMID 15223038.
  19. ^ Grazziotin, Felipe G; Zaher, Hussam; Murphy, Robert W; Scrocchi, Gustavo; Benavides, Marco A; Zhang, Ya-Ping; Bonatto, Sandro L (2012). "Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): A reappraisal". Cladistics. 28 (5): 437–459. doi:10.1111/j.1096-0031.2012.00393.x. PMID 34836446. S2CID 84934386.
  20. ^ D. A. Rossman (1965). Comments on the Revival of the Colubrid Snake Subfamily Heterodontinae.
  21. ^ He M; Feng JC; Liu SY; Guo P; Zhao EM (2009). "The phylogenetic position of Thermophis (Serpentes: Colubridae), an endemic snake from the Qinghai‐Xizang Plateau, China" (PDF). Journal of Natural History. 43 (7–8): 479–488. doi:10.1080/00222930802389825. S2CID 84653966. Archived from the original (PDF) on 2015-12-22. Retrieved 2018-05-14.
  22. ^ Huang S; Liu SY; Guo P; Zhang YP; Zhao EM (2009). "What are the closest relatives of the hot-spring snakes (Colubridae, Thermophis), the relict species endemic to the Tibetan Plateau?" (PDF). Molecular Phylogenetics and Evolution. 51 (3): 438–446. doi:10.1016/j.ympev.2009.02.013. PMID 19249375. Archived from the original (PDF) on 2013-10-03. Retrieved 2018-05-14.