Colubridae
{{Short description|Family of snakes}}
{{Automatic taxobox
| fossil_range = {{fossil range|Late Eocene|recent|Late Eocene—present}}
| image = Colubridae-01.jpg
| image_caption = Colubridae species
| taxon = Colubridae
| authority = Oppel, 1811
| subdivision_ranks =
| subdivision =
}}
Colubridae ({{IPAc-en|k|ə|ˈ|l|uː|b|r|ɪ|d|iː}}, commonly known as colubrids {{IPAc-en|ˈ|k|ɒ|lj|ʊ|b|r|ɪ|d|z}}, from {{langx|la|coluber}}, 'snake') is a family of snakes. With 249 genera,{{cite web |title=Colubrid |url=https://www.britannica.com/animal/colubrid |website=britannica.com |publisher=Britannica |access-date=2 December 2018 |archive-date=3 December 2018 |archive-url=https://web.archive.org/web/20181203013937/https://www.britannica.com/animal/colubrid |url-status=live}} it is the largest snake family. The earliest fossil species of the family date back to the Late Eocene epoch, with earlier origins suspected.{{Cite book |last1=P Dennis |url=http://archive.org/details/actazoolcrac-v461-01 |title=Nebraskophis HOLMAN from the Late Eocene of Georgia (USA), the oldest known North American colubrid snake |last2=A J |date=2003}} Colubrid snakes are found on every continent except Antarctica.{{cite book |editor=Cogger, H.G.|editor-link=Harold Cogger |editor2=Zweifel, R.G.|editor2-link=Richard G. Zweifel |author=Bauer, Aaron M. |year=1998 |title=Encyclopedia of Reptiles and Amphibians |publisher=Academic Press |location=San Diego |pages=188–195 |isbn=978-0-12-178560-4 |author-link=:fr:Aaron Matthew Bauer}}
Description
Colubrids are a very diverse group of snakes. They can exhibit many different body styles, body sizes, colors, and patterns. They can also live in many different types of habitats including aquatic, terrestrial, semi-arboreal, arboreal, desert, mountainous forests, semi-fossorial, and brackish waters.{{cite book|last1=Vitt|first1=Laurie J.|first2=Janalee P.|last2=Caldwell|title=Herpetology: An Introductory Biology of Amphibians and Reptiles|edition=4th|publisher=Elsevier Inc|year=2013|isbn=978-0-12-386919-7|pages=622–626}}{{rp|622-623}}
A primarily shy and harmless group of snakes, the vast majority of colubrids are not venomous, nor do most colubrids produce venom that is medically significant to mammals. However, the bites of some can escalate quickly to emergency situations. Furthermore, within the Colubridae, the South African boomslang and twig snakes, as well as the Asian keelback snakes (Rhabdophis sp.) have long been notorious for inflicting the worst bites on humans, with the most confirmed fatalities.{{cite journal |author=Bruna Azara, C. |year=1995 |title=Animales venenosos. Vertebrados terrestres venenosos peligrosos para el ser humano en España |url=http://www.sea-entomologia.org/PDF/BOLETIN_11/B11-009-032.pdf |journal=Boletín de la S.E.A. |volume=11 |pages=32–40 |access-date=2016-09-30 |archive-date=2020-08-14 |archive-url=https://web.archive.org/web/20200814031303/http://sea-entomologia.org/PDF/BOLETIN_11/B11-009-032.pdf |url-status=live}}{{cite book |doi=10.1016/C2010-0-68461-6 |title="Venomous" Bites from Non-Venomous Snakes |date=2011 |isbn=978-0-12-387732-1 }}
Some colubrids are described as opisthoglyphous (often simply called "rear-fanged"), meaning they possess shortened, grooved "fangs" located at the back of the upper jaw. It is thought that opisthoglyphy evolved many times throughout the natural history of squamates and is an evolutionary precursor to the larger, frontal fangs of vipers and elapids.{{cite journal |last1=Jackson |first1=K |title=The evolution of venom-delivery systems in snakes |journal=Zoological Journal of the Linnean Society |date=2003 |volume=137 |issue=3 |pages=337–354 |doi=10.1046/j.1096-3642.2003.00052.x |doi-access=free }}{{cite journal |last1=Vonk |first1=F. J. |last2=Admiraal |first2=J. F. |last3=Jackson |first3=K. |last4=Reshef |first4=R. |last5=de Bakker |first5=M. A. |last6=Vanderschoot |first6=K. |last7=van den Berge |first7=I. |last8=van Atten |first8=M. |last9=Burgerhout |first9=E.|last10=Beck|first10=A. |title=Evolutionary origin and development of snake fangs |journal=Nature |date=2008 |volume=454 |issue=7204 |pages=630–633 |doi=10.1038/nature07178 |pmid=18668106 |bibcode=2008Natur.454..630V |s2cid=4362616 }}{{cite journal |last1=Fry |first1=B. G. |last2=Casewell |first2=N. R. |last3=Wüster |first3=W. |last4=Vidal |first4=N. |last5=Young |first5=B. |last6=Jackson |first6=T. N. |title=The structural and functional diversification of the Toxicofera reptile venom system |journal=Toxicon |date=2012 |volume=60 |issue=4 |pages=434–448 |doi=10.1016/j.toxicon.2012.02.013 |pmid=22446061 |bibcode=2012Txcn...60..434F }} These grooved fangs tend to be sharpest on the anterior and posterior edges.{{Cite journal |last1=Cleuren |first1=Silke G. C. |last2=Hocking |first2=David P. |last3=Evans |first3=Alistair R. |date=June 2021 |title=Fang evolution in venomous snakes: Adaptation of 3D tooth shape to the biomechanical properties of their prey |journal=Evolution |language=en |volume=75 |issue=6 |pages=1377–1394 |doi=10.1111/evo.14239 |pmid=33904594 |url=https://research.monash.edu/en/publications/334aa3e9-42d2-4947-8bd4-f8cb8e6b75d0 }} While feeding, colubrids move their jaws backward to create a cutting motion between the posterior edge and the prey's tissue. In order to inject venom, colubridae must chew on their prey.{{cite book |doi=10.1016/B978-0-12-824315-2.00652-7 |chapter=Snakes |title=Encyclopedia of Toxicology |date=2024 |last1=Schmalzried |first1=Scott |last2=Ceretto |first2=Vincent |pages=567–571 |isbn=978-0-323-85434-4 }} Colubrids can also be proteroglyphous (fangs at the front of the upper jaw, followed by small solid teeth)
Most Colubridae are oviparous (mode of reproduction where an egg is produced that will later hatch) with clutch size varying by size and species of snake. However, certain species of snakes from the subfamilies of Natricinae and Colubrinae are viviparous (mode of reproduction where young are live birthed). These viviparous species can birth various amounts of offspring at a time, but the exact number of offspring depends on the size and species of snake.
Characteristics of Colubridae
Characteristics of Colubridae include limbless bodies, left lung that is reduced or absent with or without a tracheal lung, well-developed oviducts, premaxillaries that lack teeth, maxilaries oriented longitudinally with teeth that are solid or grooved, mandible without a coronoid bone, dentary that has teeth, only a left carotid artery, intracostal arteries arising from the dorsal aorta every few trunk segments, no cranial infrared receptors occurring in pits or surface indentations, and optic foramina that typically traverse the frontal–parietal–parasphenoid sutures.
Classification
In the past, the Colubridae were not a natural group, as many were more closely related to other groups, such as elapids, than to each other.{{cite journal |last1=Lawson |first1=R. |last2=Slowinski |first2=J.B. |last3=Crother |first3=B.I. |last4=Burbrink |first4=F.T. |title=Phylogeny of the Colubroidea (Serpentes): New evidence from mitochondrial and nuclear genes |journal=Molecular Phylogenetics and Evolution |date=November 2005 |volume=37 |issue=2 |pages=581–601 |doi=10.1016/j.ympev.2005.07.016 |pmid=16172004 |bibcode=2005MolPE..37..581L }} This family was historically used as a "wastebasket taxon" for snakes that do not fit elsewhere.{{cite journal |author1=Fry, B.G. |author2=Vidal, N. |author3=van der Weerd, L. |author4=Kochva, E. |author5=Renjifo, C. |title=Evolution and diversification of the Toxicofera reptile venom system |year=2009 |journal=Journal of Proteomics |volume=72 |issue=2 |pages=127–136 |doi=10.1016/j.jprot.2009.01.009 |pmid=19457354}} Until recently, colubrids were basically colubroids that were not elapids, viperids, or Atractaspis.{{cite book |last1=Pough |first1=F. Harvey |last2=Andrews |first2=R. M. |last3=Cadle |first3=J. E. |last4=Crump |first4=M. L. |last5=Savitzky |first5=A. H. |last6=Wells |first6=K. |title=Herpetology |date=1998 |publisher=Prentice Hall |isbn=978-0-13-850876-0 |page=162}}
However, recent research in molecular phylogenetics has stabilized the classification of historically "colubrid" snakes and the family as currently defined is a monophyletic clade,{{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 |issue=1 |pages=93 |doi=10.1186/1471-2148-13-93 |pmid=23627680 |pmc=3682911 |bibcode=2013BMCEE..13...93P |doi-access=free }}{{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 |pmc=5014348 |pmid=27603205 |doi=10.1371/journal.pone.0161070 |bibcode=2016PLoSO..1161070F |doi-access=free}}{{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 |pmid=21074626 |doi=10.1016/j.ympev.2010.11.006 |bibcode=2011MolPE..58..329P }}{{cite journal |last1=Zheng |first1=Yuchi |last2=Wiens |first2=John J. |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=January 2016 |volume=94 |issue=Pt B |pages=537–547 |pmid=26475614 |doi=10.1016/j.ympev.2015.10.009 |bibcode=2016MolPE..94..537Z }} although additional research will be necessary to sort out all the relationships within this group. As of May 2018, eight subfamilies are recognized.{{cite web |last1=Uetz |first1=Peter |title=Colubridae at The Reptile Database |url=http://reptile-database.reptarium.cz/advanced_search?taxon=Colubridae&submit=Search |website=The Reptile Database |publisher=EMBL |access-date=13 May 2018 |archive-date=29 June 2018 |archive-url=https://web.archive.org/web/20180629172949/http://reptile-database.reptarium.cz/advanced_search?taxon=Colubridae&submit=Search |url-status=live}}
=Current subfamilies=
Sibynophiinae – three genera
{{columns-list|colwidth=15em|
}}
Natricinae – 36 genera (sometimes given as family Natricidae)
Image:Indian Rat Snake (Grey and Yellow).jpg (grey and yellow)]]
Image:Coast_Garter_Snake.jpg, Thamnophis sirtalis]]
{{columns-list|colwidth=15em|
- Afronatrix
- Amphiesma
- Amphiesmoides
- Anoplohydrus
- Aspidura
- Atretium
- Blythia{{Cite web |url=https://reptile-database.reptarium.cz/species?genus=Blythia&species=reticulata&search_param=%28%28genus%3D%27Blythia%27%29%29 |title=Blythia reticulata |access-date=2021-05-21 |archive-date=2021-05-21 |archive-url=https://web.archive.org/web/20210521033938/https://reptile-database.reptarium.cz/species?genus=Blythia&species=reticulata&search_param=%28%28genus%3D%27Blythia%27%29%29 |url-status=live}}
- Clonophis
- Fowlea
- Haldea
- Hebius
- Helophis
- Herpetoreas
- Hydrablabes
- Hydraethiops
- Iguanognathus
- Isanophis
- Limnophis
- Liodytes
- Natriciteres
- Natrix
- †Neonatrix
- Nerodia
- Opisthotropis
- Pseudagkistrodon
- Regina
- Rhabdophis
- Rhabdops
- Smithophis
- Storeria
- Thamnophis
- Trachischium
- Trimerodytes
- Tropidoclonion
- Tropidonophis
- Virginia
- Xenochrophis}}
Pseudoxenodontinae – two genera
{{columns-list|colwidth=15em|
}}
Dipsadinae – over 100 genera (sometimes given as family Dipsadidae)
Image:Sibon_longifrenis_(La_Selva_Biological_Station).jpg, Sibon longifrenis]]
{{columns-list|colwidth=15em|
- Adelphicos
- Adelphostigma
- Alsophis
- Amastridium
- Amnesteophis
- Amnisiophis
- Apographon
- Apostolepis
- Arcanumophis
- Arrhyton
- Atractus
- Baliodryas
- Boiruna
- Borikenophis
- Caaeteboia
- Calamodontophis
- Caraiba
- Carphophis
- Cenaspis
- Cercophis
- Chersodromus
- Chlorosoma
- Clelia
- Coniophanes
- Conophis
- Contia
- Coronelaps
- Crisantophis
- Cryophis
- Cubophis
- Diadophis
- Diaphorolepis
- Dibernardia
- Dipsas
- Ditaxodon
- Drepanoides
- Dryophylax
- Echinanthera
- Elapomorphus
- Emmochliophis
- Enuliophis
- Enulius
- Erythrolamprus
- Eutrachelophis
- Farancia
- Galvarinus
- Geophis
- Gomesophis
- Haitiophis
- Helicops
- Heterodon
- Hydrodynastes
- Hydromorphus
- Hydrops
- Hypsiglena
- Hypsirhynchus
- Ialtris
- Imantodes
- Incaspis
- Leptodeira
- Lioheterophis
- Lygophis
- Magliophis
- Manolepis
- Mesotes
- Mussurana
- Ninia
- Nothopsis
- Omoadiphas
- Oxyrhopus
- Paikwaophis
- †Paleoheterodon
- Paraphimophis
- Phalotris
- Philodryas
- Phimophis
- Plesiodipsas
- Pliocercus
- Pseudalsophis
- Pseudoboa
- Pseudoeryx
- Pseudoleptodeira
- Psomophis
- Ptychophis
- Rhachidelus
- Rhadinaea
- Rhadinella
- Rhadinophanes
- Rodriguesophis
- Saphenophis
- Sibon
- Siphlophis
- Sordellina
- Synophis
- Tachymenis
- Tachymenoides
- Taeniophallus
- Tantalophis
- Thamnodynastes
- Thermophis
- Tomodon
- Tretanorhinus
- Trimetopon
- Tropidodipsas
- Tropidodryas
- Uromacer
- Urotheca
- Xenodon
- Xenopholis
- Xenoxybelis
- Zonateres
}}
Grayiinae – one genus
Calamariinae – seven genera
{{columns-list|colwidth=15em|
}}
Ahaetuliinae – five genera
{{columns-list|colwidth=15em|
Colubrinae – 93 genera
Image:PikiWiki Israel 37648 Nature and Colors.jpg, Dolichophis jugularis, preying on a legless lizard, a sheltopusik]]
{{columns-list|colwidth=15em|
- Aeluroglena
- Aprosdoketophis
- Archelaphe
- Arizona
- Bamanophis
- Bogertophis
- Boiga
- Cemophora
- Chapinophis
- Chironius
- Coelognathus
- Coluber
- Conopsis
- Coronella
- Crotaphopeltis
- Dasypeltis
- Dendrophidion
- Dipsadoboa
- Dispholidus
- Dolichophis
- Drymarchon
- Drymobius
- Drymoluber
- Eirenis
- Elaphe
- Euprepiophis
- Ficimia
- Geagras
- Gonyosoma
- Gyalopion
- Hapsidophrys
- Hemerophis
- Hemorrhois
- Hierophis
- Lampropeltis
- Leptodrymus
- Leptophis
- Liopeltis
- Lycodon
- Lytorhynchus
- Macroprotodon
- Masticophis
- Mastigodryas
- Meizodon
- Mopanveldophis
- Muhtarophis
- Oligodon
- Oocatochus
- Opheodrys
- Oreocryptophis
- Orientocoluber
- Oxybelis
- Palusophis
- Pantherophis
- {{extinct}}Paracoluber
- Persiophis
- Philothamnus
- Phrynonax
- Phyllorhynchus
- Pituophis
- Platyceps
- Pseudelaphe
- Pseudoficimia
- Ptyas
- Rhamnophis
- Rhinobothryum
- Rhinocheilus
- Rhynchocalamus
- Salvadora
- Scaphiophis
- Scolecophis
- Senticolis
- Simophis
- Sonora
- Spalerosophis
- Spilotes
- Stegonotus
- Stenorrhina
- Stichophanes
- Symphimus
- Sympholis
- Tantilla
- Tantillita
- Telescopus
- Thelotornis
- Thrasops
- Toxicodryas
- Trimorphodon
- Wallaceophis
- Wallophis
- Xenelaphis
- Xyelodontophis
- Zamenis
- †Zelceophis}}
Sub-family currently undetermined
{{columns-list|colwidth=15em|
- Elapoidis
- Gongylosoma
- Lycognathophis
- Oreocalamus
- Tetralepis
- †Ameiseophis{{Cite journal |last=Holman |first=J. Alan |date=1982-01-01 |title=New Herpetological Species and Records from the Norden Bridge Fauna (Miocene: Late Barstovian) of Nebraska |journal=Transactions of the Nebraska Academy of Sciences and Affiliated Societies |url=https://digitalcommons.unl.edu/tnas/491}}
- †Dryinoides
- †Hispanophis
- †Floridaophis{{Cite web |title=Floridaophis auffenbergi |url=https://www.floridamuseum.ufl.edu/florida-vertebrate-fossils/species/floridaophis-auffenbergi/ |access-date=2023-12-15 |website=Florida Vertebrate Fossils |language=en-US}}
- †Micronatrix
- †Miocoluber
- †Mionatrix?
- †Nebraskophis
- †Palaeonatrix
- †Paleofarancia
- †Paraoxybelis
- †Paraxenophis
- †Periergophis
- †Pollackophis
- †Pseudocemophora
- †Texasophis}}
=Former subfamilies=
These taxa have been at one time or another classified as part of the Colubridae, but are now either classified as parts of other families, or are no longer accepted because all the species within them have been moved to other (sub)families.
- Subfamily Aparallactinae (now a subfamily of Lamprophiidae, sometimes combined with Atractaspidinae)
- Subfamily Boiginae (now part of Colubrinae)
- Subfamily Boodontinae (some of which now treated as subfamily Grayiinae of the new Colubridae, others moved to family Lamprophiidae as part of subfamilies Lamprophiinae, Pseudaspidinae and Pseudoxyrhophiidae, which are now sometimes treated as families)
- Subfamily Dispholidinae (now part of Colubrinae)
- Subfamily Homalopsinae (now family Homalopsidae)
- Subfamily Lamprophiinae (now a subfamily of Lamprophiidae)
- Subfamily Lycodontinae (now part of Colubrinae)
- Subfamily Lycophidinae (now part of Lamprophiidae)
- Subfamily Pareatinae (now family Pareidae, sometimes incorrectly spelled Pareatidae){{cite journal |last1=Savage |first1=Jay M. |year=2015 |title=What are the correct family names for the taxa that include the snake genera Xenodermus, Pareas, and Calamaria? |journal=Herpetological Review |volume=46 |issue=4 |pages=664–665 |url=http://www.zenscientist.com/index.php/pdflibrary2/func-finishdown/2496/ |access-date=2018-05-14 |archive-url=https://web.archive.org/web/20160307141006/http://www.zenscientist.com/index.php/pdflibrary2/func-finishdown/2496 |archive-date=2016-03-07 |url-status=dead}}
- Subfamily Philothamninae (now part of Colubrinae)
- Subfamily Psammophiinae (now a subfamily of Lamprophiidae)
- Subfamily Pseudoxyrhophiinae (now a subfamily of Lamprophiidae)
- Subfamily Xenoderminae (now family Xenodermidae, sometimes incorrectly spelled Xenodermatidae)
- Subfamily Xenodontinae (which many authors put in Dipsadinae/Dipsadidae)
Fossil record
The oldest colubrid fossils are indeterminate vertebrae from Thailand and specimens of the genus Nebraskophis from the U.S. state of Georgia, both from the Late Eocene. The presence of derived colubrids in North America so early on, despite their presumed Old World origins, suggests that they originated even earlier. The Pliocene (Blancan) fossil record in the Ringold Formation of Adams County, Washington has yielded fossils from a number of colubrids including Elaphe pliocenica, Elaphe vulpina, Lampropeltis getulus, Pituophis catenifer, a Thamnophis species, and the extinct genus Tauntonophis.{{cite journal |last1=Parmley |first1=D. |last2=Walker |first2=D. |year=2003 |title=Snakes of the Pliocene Taunton local fauna of Adams County, Washington with the description of a new colubrid |journal=Journal of Herpetology |volume=37 |issue=2 |pages=235–244 |doi=10.1670/0022-1511(2003)037[0235:SOTPTL]2.0.CO;2|s2cid=86000331}}
References
= Citations =
{{Reflist|30em}}
=Bibliography=
- {{cite journal |last1=Barbière |first1=Franck |last2=Ortiz |first2=Pablo E. |last3=Pardiñas |first3=Ulyses F.J. |title=The oldest sigmodontine rodent revisited and the age of the first South American cricetids |journal=Journal of Paleontology |date=March 2019 |volume=93 |issue=2 |pages=368–384 |doi=10.1017/jpa.2018.74 |bibcode=2019JPal...93..368B |s2cid=135378126 }}
- {{cite journal |last1=Barbiere |first1=Franck |last2=Cruz |first2=Laura E. |last3=Ortiz |first3=Pablo E. |last4=Pardiñas |first4=Ulyses F. J. |title=A new genus of Sigmodontinae (Mammalia, Rodentia, Cricetidae) from the Pliocene of central Argentina |journal=Journal of Vertebrate Paleontology |date=2 September 2016 |volume=36 |issue=5 |pages=e1199557 |doi=10.1080/02724634.2016.1199557 |bibcode=2016JVPal..36E9557B |s2cid=89495030 |hdl=11336/36864 |hdl-access=free }}
- {{cite journal |last1=Nicoli |first1=Laura |title=New fossil species of the extant genus Lepidobatrachus (Anura, Ceratophryidae) from the Late Miocene-Early Pliocene of central Argentina |journal=Journal of Vertebrate Paleontology |date=3 September 2015 |volume=35 |issue=5 |pages=e981636 |doi=10.1080/02724634.2015.981636 |bibcode=2015JVPal..35E1636N |hdl=11336/46258 |s2cid=83574417 |hdl-access=free }}
- {{cite journal |last1=Agnolin |first1=Federico |last2=Bogan |first2=Sergio |last3=Tomassini |first3=Rodrigo |last4=Manera |first4=Teresa |title=New Percichthyidae (Teleostei, Percoidei) from the early Pliocene of the Buenos Aires province (Argentina) and its biogeographical implications |journal=Revista del Museo Argentino de Ciencias Naturales |date=2014 |volume=16 |pages=19–31 |doi=10.22179/REVMACN.16.159 |doi-access=free }}
- {{cite journal |last1=Zamorano |first1=Martín |title=Diagnosis y nueva descripción de Propanochthus bullifer (Burmeister) (Xenarthra, Glyptodontidae). Consideraciones bioestratigráficas y cronológicas de su procedencia |journal=Spanish Journal of Palaeontology |date=20 July 2020 |volume=28 |issue=2 |pages=283–292 |doi=10.7203/sjp.28.2.17860 |url=http://sedici.unlp.edu.ar/bitstream/handle/10915/126207/Documento.pdf?sequence=1 }}
- {{cite book |last1=Vizcaíno |first1=Sergio F. |last2=Kay |first2=Richard F. |last3=Bargo |first3=M. Susana |title=Early Miocene Paleobiology in Patagonia: High-Latitude Paleocommunities of the Santa Cruz Formation |date=2012 |publisher=Cambridge University Press |isbn=978-0-521-19461-7 }}
- {{cite journal |last1=Albino |first1=Adriana M. |last2=Tomassini |first2=Rodrigo |last3=Brizuela |first3=Santiago |title=Presencia del lagarto teiido Tupinambis en la Formación Monte Hermoso de Farola Monte Hermoso, sur de la provincia de Buenos Aires (Argentina) |journal=Ameghiniana |date=2009 |volume=46 |issue=1 |pages=177–188 |url=https://ameghiniana.org.ar/index.php/ameghiniana/article/view/182 |language=es }}
- {{cite journal |last1=Bogan |first1=Sergio |last2=Agnolin |first2=Federico |title=First fossil record for the family Trichomycteridae (Teleostei: Siluriformes; Pliocene) in the Monte Hermoso Formation, Argentina |journal=Revista del Museo Argentino de Ciencias Naturales |date=2009 |volume=11 |pages=193–198 |doi=10.22179/REVMACN.11.259 }}
- {{cite journal |last1=Verzi |first1=Diego H. |last2=Montalvo |first2=Claudia I. |last3=Deschamps |first3=Cecilia M. |title=Biostratigraphy and biochronology of the Late Miocene of central Argentina: Evidence from rodents and taphonomy |journal=Geobios |date=January 2008 |volume=41 |issue=1 |pages=145–155 |doi=10.1016/j.geobios.2006.09.005 |bibcode=2008Geobi..41..145V }}
- {{cite journal |last1=Cozzuol |first1=Mario Alberto |title=The Acre vertebrate fauna: Age, diversity, and geography |journal=Journal of South American Earth Sciences |date=July 2006 |volume=21 |issue=3 |pages=185–203 |doi=10.1016/j.jsames.2006.03.005 |bibcode=2006JSAES..21..185C }}
- {{cite journal |last1=Marshall |first1=Larry G. |last2=Sempere |first2=Thierry |year=1991 |title=The Eocene to Pleistocene vertebrates of Bolivia and their stratigraphic context: A review |journal=Revista Tecnica de Yacimientos Petroliferos Fiscales Bolivianos |volume=12 |pages=631–652 |oclc=713096859 |url=https://www.documentation.ird.fr/hor/fdi:010024181 }}
External links
- [https://snakesarelong.blogspot.com/search?q=Psammophids Psammophids] at Life Is Short, but Snakes Are Long
{{Snake_families}}
{{Taxonbar|from=Q182751}}
{{Authority control}}
Category:Extant Oligocene first appearances