Viperinae

True vipers
Asp viper, Vipera aspis
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Subphylum:	Vertebrata
Class:	Reptilia
Order:	Squamata
Suborder:	Serpentes
Family:	Viperidae
Subfamily:	Viperinae Oppel, 1811
Synonyms
Viperini - Oppel, 1811 Viperes - Cuvier, 1817 Viperides - Latreille, 1825 Viperina - Gray, 1825 Viperiodea - Fitzinger, 1826 Viperiodei - Eichwald, 1831 Viperinae - Cantor, 1847 Viperiformes - Günther, 1864 Viperida - Strauch, 1869 Atherini - Broadley, 1996[1]

Common names: pitless vipers, true vipers, Old World vipers,^[2] true adders.^[3]

The Viperinae, or viperines, are a subfamily of venomous vipers found in Europe, Asia and Africa. They are distinguished by their lack of the heat-sensing pit organs that characterize their sister group, the Crotalinae. Currently, 12 genera and 66 species are recognized.^[4] Most are tropical and subtropical, although one species, Vipera berus, even occurs within the Arctic Circle.^[2]

Description

Members of this subfamily range in size from Bitis schneideri, that grows to a maximum of 28 cm, to Bitis gabonica that reaches a maximum length of over 2 m. Most species are terrestrial, but a few, such as Atheris, are completely arboreal.^[2]

Although the heat-sensing pits that characterize the Crotalinae are clearly lacking in the viperines, a supernasal sac with sensory function has been described in a number of species. This sac is an invagination of the skin between the supranasal and nasal scales and is connected to the ophthalmic branch of the trigeminal nerve. The nerve endings here resemble those in the labial pits of boas. The supernasal sac is present in Daboia, Pseudocerastes and Causus, but is especially well developed in Bitis. Experiments have shown that strikes are not only guided by visual and chemical cues, but also by heat, with warmer targets being struck more frequently than colder ones.^[2]

Geographic range

Europe, Asia and Africa.^[1] However, they do not occur in Madagascar.^[5]

Reproduction

Generally, members of this subfamily are viviparous (ovoviviparous), although a few, such as Pseudocerastes, lay eggs.^[2]

Genera

Genus[4]	Taxon author[4]	Species[4]	Subsp.*[4]	Common name[2][6]	Geographic range[1]
Adenorhinos	Loveridge, 1930	1	0	Uzungwe viper	Central Tanzania: Udzungwe and Ukinga Mountains.
Atheris	Cope, 1862	8	1	Bush vipers	Tropical subsaharan Africa, excluding southern Africa.
Bitis	Gray, 1842	14	2	Puff adders	Africa and the southern Arabian Peninsula.
Cerastes	Laurenti, 1768	3	0	Horned vipers	North Africa eastward through Arabia and Iran.
Daboia	Gray, 1842	1	1	Russell's viper	Pakistan, India, Sri Lanka, Bangladesh, Nepal, Myanmar, Thailand, Cambodia, China (Kwangsi and Kwangtung), Taiwan and Indonesia (Endeh, Flores, east Java, Komodo, Lomblen Islands).
Echis	Merrem, 1820	8	6	Saw-scaled vipers	India and Sri Lanka, parts of the Middle East and Africa north of the equator.
Eristicophis	Alcock and Finn, 1897	1	0	McMahon's viper	The desert region of Balochistan near the Iran-Afghanistan-Pakistan border.
Macrovipera	Reuss, 1927	4	4	Large Palearctic vipers	Semideserts and steppes of northern Africa, the Near and Middle East, and the Milos Archipelago in the Aegean Sea.
Montatheris	Boulenger, 1910	1	0	Kenya mountain viper	Kenya: moorlands of the Aberdare range and Mount Kenya above 3000 m.
Proatheris	Peters, 1854	1	0	Lowland viper	Floodplains from southern Tanzania (northern end of Lake Malawi) through Malawi to near Beira, central Mozambique.
Pseudocerastes	Boulenger, 1896	1	1	False horned viper	From the Sinai of Egypt eastward to Pakistan.
ViperaT	Laurenti, 1768	23	12	Palearctic vipers	Great Britain and nearly all of continental Europe across the Arctic Circle and on some islands in the Mediterranean (Elba, Montecristo, Sicily) and Aegean Sea eastward across northern Asia to Sakhalin Island and North Korea. Also found in northern Africa in Morocco, Algeria and Tunisia.

*) Not including the nominate subspecies.
^T) Type genus.

Taxonomy

Until relatively recently, two other genera were also included in the Viperinae. However, they were eventually considered so distinctive within the Viperidae, that separate subfamilies were created for them:^[1]

• Genus Azemiops - moved to subfamily Azemiopinae by Liem, Marx & Rabb (1971).
• Genus Causus - recognition of subfamily Causinae (Cope, 1860) was proposed by Groombridge (1987) and further supported by Cadle (1992).

Nevertheless, these groups, together with the genera currently recognized as belonging to the Viperinae, are still often referred to collectively as the true vipers.^[2]

Broadley (1996) recognized a new tribe, Atherini, for the genera Atheris, Adenorhinos, Montatheris and Proatheris, the type genus for which is Atheris.^[1]

See also

• List of viperine species and subspecies
• Viperinae by common name
• Viperinae by taxonomic synonyms
• Snakebite

References

1. ^ ^{a b c d e} McDiarmid RW, Campbell JA, Touré T. 1999. Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1. Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
2. ^ ^{a b c d e f g} Mallow D, Ludwig D, Nilson G. 2003. True Vipers: Natural History and Toxinology of Old World Vipers. Krieger Publishing Company, Malabar, Florida. 359 pp. ISBN 0-89464-877-2.
3. ^ U.S. Navy. 1991. Poisonous Snakes of the World. US Govt. New York: Dover Publications Inc. 203 pp. ISBN 0-486-26629-X.
4. ^ ^{a b c d e} "Viperinae". Integrated Taxonomic Information System. http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=563898. Retrieved 4 August 2006. 
5. ^ Stidworthy J. 1974. Snakes of the World. Grosset & Dunlap Inc. 160 pp. ISBN 0-448-11856-4.
6. ^ Spawls S, Branch B. 1995. The Dangerous Snakes of Africa. Ralph Curtis Books. Dubai: Oriental Press. 192 pp. ISBN 0-88359-029-8.

Further reading

• Breidenbach CH. 1990. Thermal cues influence strikes in pitless vipers. Journal of Herpetology 4: 448-50.
• Broadley DG. 1996. A review of the tribe Atherini (Serpentes: Viperidae), with the descriptions of two new genera. African Journal of Herpetology 45(2): 40-48.
• Cantor TE. 1847. Catalogue of reptiles inhabiting the Malayan Peninsula and Islands. Journal of the Asiatic Society of Bengal. Calcutta. 16(2): 607-656, 897-952, 1026-1078[1040].
• Cuvier G. 1817. Le règne animal distribué d'après son organisation, pour servir de base à l'histoire naturelle des animaux det d'introduction à l'anatomie comparée. Tome II, contenant les reptiles, les poissons, les mollusques et les annélidés. Déterville, Paris. xviii, 532 pp.[80].
• Eichwald, E. 1831. Zoologia specialis, quam expositis animalibus tum vivis, tum fossilibus potissimuni rossiae in universum, et poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi. Zawadski, Vilnae. 3: 404 pp.[371].
• Fitzinger LJFJ. 1826. Neue classification der reptilien nach ihren natürlichen verwandtschaften. Nebst einer verwandtschafts-tafel und einem verzeichnisse der reptilien-sammlung des K. K. zoologischen museum's zu Wien. J.G. Hübner, Wien. vii, 66 pp.[11].
• Gray JE. 1825. A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy, new ser., 10: 193-217[205].
• Günther ACLG. 1864. The Reptiles of British India. Ray Society. London. xxvii. 452 pp.[383].
• Latreille PA. 1825. Familles naturelles du règne animal, exposés succinctement et dans un ordre analytique, avec l'indication de leurs genres. Bailliere, Paris. 570 pp.[102].
• Lynn WG. 1931. The structure and function of the facial pit of the pit vipers. American Journal of Anatomy 49: 97.
• Oppel M. 1811. Mémoire sur la classification des reptiles. Ordre II. Reptiles à écailles. Section II. Ophidiens. Annales du Musée National d'Histoire Naturelle, Paris 16: 254-295, 376-393.[376, 378, 389].
• Strauch A. 1869. Mémoires de l'Academie des Sciences Impérial de St. Pétersbourg (7)14: 144 pp.[19]