Viperinae

Viperinae
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

Viperinae is a subfamily of terrestrial and arboreal venomous vipers (family Viperidae) characterized by a lack of the heat-sensing pit organs that characterize their sister group, the subfamily Crotalinae. Viperines are found in Europe, Asia, and Africa, and while largely tropical and subtropical, one species even occurs within the Arctic Circle. Technically known as viperines, members of Viperinae also are known by the common names of true vipers, pitless vipers, Old World vipers, or true adders. Well known members include the puff adders and bush vipers. About 66 species in 12 genera are recognized.

Because of their venom, viperines are highly feared by people. The African puff adder (Bitis arietans) has a wide distribution in Africa, is relatively common, has large size and potent venom, and exhibits a willingness to bite, all make it responsible for more fatalities than any other African snake. The saw-scaled viper (Echis carinatus) and Russell's viper (Daboia sp.), found in Asia, likewise have a powerful venom and cause a number of deaths.

Nonetheless, viperines also provide an important ecological role, helping to control populations of prey species such as rodents, scorpions, squirrels, centipedes, and so forth. They also provide food for some animals, such as birds of prey. For humans, not only do they add to the wonder of nature, but the venom offers great potential for medical use because of the specificity of the compounds on the body; some venoms, for example, are used to treat blood disorders. Viperines also have been used in Africa for a crude form of hunting, whereby a captured viperine is placed on a game trail to strike an animal using the trail.

Overview and description

Viperidae, the taxon to which Viperinae belongs, is a family of venomous snakes characterized by a head that is distinct from the body and by a pair of long, hollow, venom-injecting fangs that can be folded back against the top of the mouth (tip inward) when the mouth is closed. The two main subfamilies in Viperidae are Viperinae (pitless vipers or true vipers) and Crotalinae (pit vipers), although other subfamilies also are recognized. Members of the family Viperidae commonly are known as vipers. However, the term viperid also is used for members of Viperidae. Viperid offers greater specificity than the term viper since some authorities use "viper" for members of the subfamily Viperinae and "pit viper" for members of the subfamily Crotalinae (Nilson and Gutverlet 2004).

Technically, members of Viperinae may be known as viperines. However, they also are known commonly as pitless vipers, true vipers, Old World vipers (Mallow et al. 2003), and true adders (U.S. Navy 1991). Currently, 12 genera and 66 species are recognized (ITIS 2004).

Viperines differ from their sister group, the Crotalinae, by the lack of a heat-sensing pit organ, which in the pit vipers are located between the eye and the nostril on either side of the head. Members of the Viperinae subfamily range in size from Bitis schneideri, which grows to a maximum of 28 centimeters (11 inches), to Bitis gabonica, which reaches a maximum length of over 2 meters (6.5 feet).

Most species are terrestrial, but a few, such as Atheris, are completely arboreal (Mallow et al. 2003). Generally, members of this subfamily are viviparous (ovoviviparous), although a few, such as Pseudocerastes, lay eggs (Mallow et al. 2003).

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 the genera Daboia, Pseudocerastes, and Causus, but is especially well developed in Bitis. Experiments have shown that viperine strikes are not only guided by visual and chemical cues, but also by heat, with warmer targets being struck more frequently than colder ones (Mallow et al. 2003).

Geographic range

Viperinae are found in the Old World, in Europe, Asia, and Africa (McDiarmid et al. 1999). However, they do not occur in Madagascar (Stidworthy 1974).

Most are tropical and subtropical, although Vipera berus occurs within the Arctic Circle (Mallow et al. 2003). Vipera berus is extremely widespread and can be found throughout most of Western Europe and all the way to Far East Asia.

Genera

Genus	Taxon author	Species	Subsp.*	Common name	Geographic range
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 Kwantung), 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 meters.
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.

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 (McDiarmid et al. 1999):

• Genus Azemiops—moved to subfamily Azemiopinae by Liem et al. (1971).
• Genus Causus—recognition of subfamily Causinae (Cope, 1860) was proposed by B. C. Groombridge and further supported by J. E. Cadle.

Nevertheless, these groups, together with the genera currently recognized as belonging to the Viperinae, are still often referred to collectively as the true vipers (Mallow et al. 2003).

Broadley (1996) recognized a new tribe, Atherini, for the genera Atheris, Adenorhinos, Montatheris, and Proatheris, the type genus for which is Atheris (McDiarmid et al. 1999).

Further reading

• Cantor, T. E. 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].
• Gray JE. 1825. A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy 10: 193-217.
• Günther, A. C. L. G. 1864. The Reptiles of British India. London: Ray Society.
• Lynn, W. G. 1931. The structure and function of the facial pit of the pit vipers. American Journal of Anatomy 49: 97.

References

ISBN links support NWE through referral fees

• Breidenbach C. H. 1990. Thermal cues influence strikes in pitless vipers. Journal of Herpetology 4: 448-50.
• Broadley D. G. 1996. A review of the tribe Atherini (Serpentes: Viperidae), with the descriptions of two new genera. African Journal of Herpetology 45(2): 40-48.
• Integrated Taxonomic Information System (ITIS). 2004. Viperinae Oppel, 1811. ITIS Taxonomic Serial No.: 563898. Retrieved January 20, 2009.
• Liem, K. L., H. Marx, and G. R. Rabb. 1971. The viperid snake Azemiops: Its comparative cephalic anatomy and phylogenetic position in relation to Viperinae and Crotalinae. Fieldiana Zool. 59: 65– 126
• Mallow, D., D. Ludwig, and G. Nilson. 2003. True Vipers: Natural History and Toxinology of Old World Vipers. Malabar, FL: Krieger. ISBN 0894648772.
• McDiarmid, R. W., J. A. Campbell, and T. Touré. 1999. Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1. Washington, DC: Herpetologists' League. ISBN 1893777006.
• Nilson, G., and R. L. Gutberlet. 2004. Viperidae. In B. Grzimek, D. G. Kleiman, V. Geist, and M. C. McDade (eds.). Grzimek's Animal Life Encyclopedia. Detroit: Thomson-Gale. ISBN 0787657883.
• Spawls, S, and B. Branch. 1995. The Dangerous Snakes of Africa. Ralph Curtis Books. Dubai: Oriental Press. ISBN 0883590298.
• Stidworthy, J. 1974. Snakes of the World. Grosset & Dunlap. ISBN 0448118564.
• United States. Dept. of the Navy, Bureau of Medicine and Surgery. 1991. Poisonous Snakes of the World. New York: Dover Publications. ISBN 048626629X.

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