Vipera ammodytes

Vipera ammodytes
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Subphylum:	Vertebrata
Class:	Reptilia
Order:	Squamata
Suborder:	Serpentes
Family:	Viperidae
Subfamily:	Viperinae
Genus:	Vipera
Species:	V. ammodytes
Binomial name
Vipera ammodytes (Linnaeus, 1758)
Synonyms
Coluber Ammodytes - Linnaeus, 1758 Vipera Illyrica - Laurenti, 1768 Vipera ammodytes - Sonnini & Latreille, 1801 [Vipera (Echidna)] Ammodytes - Merrem, 1820 Cobra ammodytes - Fitzinger, 1826 [Pelias] Col[uber]. ammodytes - Boie, 1827 Vipera (Rhinechis) Ammodytes - Fitzinger, 1843 V[ipera]. (Vipera) ammodytes - Jan, 1863 Vipera ammodytes - Eber, 1863 Vipera ammodytes - Boulenger, 1896 [Vipera ammodytes] var. steindachneri - Werner, 1897 Vipera ammodytes [ammodytes] - Boulenger, 1903 Vipera ammodytes ammodytes - Zarevsky, 1915 Teleovipera ammodytes - Reuss, 1927 Vipera ammodytes ammodytes - Mertens & Müller, 1928 Rhinaspis illyrica litoralis - Reuss, 1935 Rhinaspis illyrica velebitensis - Reuss, 1935 Rhinaspis illyrica f[orma]. melanura - Reuss, 1937 Vipera ammodytes ruffoi - Bruno, 1968 Vipera (Rhinaspis) ammodytes ammodytes - Obst, 1983 Vipera ammodytes - Golay et al., 1993[1]

Common names: horned viper, long-nosed viper, nose-horned viper, sand viper,^[2] more.

Vipera ammodytes is a venomous viper species found in southern Europe through to the Balkans and parts of the Middle East. It is reputed to be the most dangerous of the European vipers due to its large size, long fangs (up to 13 mm) and high venom toxicity.^[3] The specific name is derived from the Greek words ammos and dutes, meaning "sand" and "burrower" or "diver"; not a very good name for an animal that actually prefers rocky habitats.^[4] Five subspecies are currently recognized, including the nominate subspecies described here.^[5]

Description

V. ammodytes

Grows to a maximum length of 95 cm (37.62 in), although individuals usually measure less than 85 cm (33.66 in). Females are somewhat smaller than males. Maximum length also depends on race, with northern forms distinctly larger than southern ones.^[3] According to Strugariu (2006), the average length is 50–70 cm (20 to 28 in) with reports of specimens over 1 m (40 in) in length. Females are usually larger and more heavily built, although the largest specimens on record are males.^[6]

The head is covered in small, irregular scales that are either smooth or only weakly keeled, except for a pair of large supraocular scales that extend beyond the posterior margin of the eye. 10-13 small scaled border the eye and two rows separate the eye from the supralabials. The nasal scale is large, single (rarely divided) and separated from the rostral by a single nasorostral scale. The rostral scale is wider than it is long.^[3]

The most distinctive characteristic is a single "horn" on the snout, just above the rostral scale. It consists of 9-17 scales arranged in 2 (rarely 2 or 4) transverse rows.^[3] It grows to a length of about 5 mm and is actually soft and flexible. In southern subspecies, the horn sits vertically upright, while in V. a. ammodytes it points diagonally forward.^[2]

The body is covered with strongly keeled dorsal scales in 21 or 23 rows (rarely 25) mid-body. The scales bordering the ventrals are smooth or weakly keeled. Males have 133-161 ventral scales and 27-46 paired subcaudals. Females have 135-164 and 24-38 respectively. The anal scale is single.^[3]

V. ammodytes

The color pattern is different for males and females. In males, the head has irregular dark brown, dark gray or black markings. A thick, black stripe runs from behind the eye to behind the angle of the jaw. The tongue is usually black and the iris has a golden or coppery color. Males have a characteristic dark blotch or V marking on the back of the head that often connects to the dorsal zigzag pattern. The ground color for males varies and includes many different shades of grey, sometimes yellowish or pinkish grey, or yellowish brown. The dorsal zigzag is dark grey or black, the edge of which is sometimes darker. A row of indistinct, dark (occasionally yellowish) spots runs along each side, sometimes joined in a wavy band.^[3]

Females have a similar color pattern, except that it is less distinct and contrasting. They usually lack the dark blotch or V marking on the back of the head that the males have. Ground color is variable and tends more towards browns and bronzes, such as grayish brown, reddish brown, copper, "dirty cream", or brick red. The dorsal zigzag is a shade of brown.^[3]

Both sexes have a zigzag dorsal stripe set against a lighter background. This pattern is often fragmented. The belly color varies and can be grayish, yellowish brown, or pinkish, "heavily clouded" with dark spots. Sometimes the ventral color is black or bluish gray with white flecks and inclusions edged in white. The chin is lighter in color than the belly. Underneath, the tip of the tail may be yellow, orange, orange-red, red or green. Melanism does occur, but is rare. Juvenile color patterns are about the same as the adults.^[3]

Common names

Horned viper, long-nosed viper, nose-horned viper, sand viper,^[2] sand adder, common sand adder, common sand viper,^[7] sand natter.^[8]

Geographic range

Southern Austria, north-eastern Italy, Slovenia, Croatia, Bosnia-Herzegovina, Serbia, Montenegro, Albania, Republic of Macedonia, Greece (including Greek Macedonia and Cyclades), Romania, Bulgaria, Turkey, Georgia and Syria. The type locality is listed as "Oriente." Schwartz (1936) proposed that the type locality be restricted to "Zara" (Zadar, Croatia).^[1]

Conservation status

This species is listed as strictly protected (Appendix II) under the Berne Convention.^[9]

Habitat

The common name sand viper is misleading, as this species does not occur in really sandy areas.^[10] Mainly, it inhabits dry, rocky hillsides with sparse vegetation. Not usually associated with woodlands, but if so it will be found there around the edges and in clearings. Sometimes found in areas of human habitation, such as railway embankments, farmland, and especially vineyards if rubble piles and stone walls are present. May be found above 2000 m at lower latitudes.^[3]

Behaviour

This species has no particular preference for its daily activity period. At higher altitudes, it is more active during the day. At lower altitudes, it may be found at any time of the day, becoming increasingly nocturnal as daytime temperatures rise.^[3]

Despite its reputation, this species is generally lethargic, not at all aggressive, and tends not to bite without considerable provocation. If surprised, wild specimens may react in a number of different ways. Some remain motionless and hiss loudly, some hiss and then flee, while still others will attempt to bite immediately.^[3]

V. ammodytes hibernates in the winter for a period of 2 to 6 months depending on environmental conditions.^[6]

V. ammodytes

Feeding

Primarily feeds on small mammals and birds. Juveniles apparently prefer lizards. Feeding behavior is influenced by prey size. Larger prey are struck, released, tracked and swallowed, while smaller prey is swallowed without using the venom apparatus. Occasionally, other snakes are eaten.^[3] There are also reports of cannibalism.^[6]

Reproduction

Before mating, the males of this species will engage an a combat dance, similar to adders.^[3] Mating takes place in the spring (April–May) and between one and twenty live young are born in August–October. At birth, juveniles are 14–24 cm long.^[6] This species is ovoviviparous.^[11]

Captivity

This species has often been kept in captivity and bred successfully.^[3] It tolerates captivity much better than other European vipers, thriving in most surroundings and usually takes food easily from the start.^[11] However, as far as handling is concerned, despite its relatively placid reputation, pinning and necking this snake can be risky, as they are relatively strong and can unexpectedly jerk free from a keeper's grasp. For close examinations, it is therefore advisable to use a clear plastic restraining tube instead.^[6]

Venom

This is likely the most dangerous snake to be found in Europe. In some areas it is at least a significant medical risk; in the past fatalities were relatively frequent in the Balkans because the peasants there had a habit of walking barefoot.^[2]

The venom can be quite toxic [based on tests conducted solely on mice], but varies over time and among different populations.^[3] Brown (1973) gives an LD₅₀ for mice of 1.2 mg/kg IV, 1.5 mg/kg IP and 2.0 mg/kg SC.^[12] Novak et al. (1973) give ranges of 0.44-0.82 mg/kg and IV and 0.19-0.64 mg/kg IP. Minton (1974) states 6.6 mg/kg SC.^[3]

The venom has both proteolytic and neurotoxic components and contains hemotoxins with blood coagulant properties, similar to and as powerful as in crotalid venom. Other properties include anticoagulant effects, hemoconcentration and hemorrhage. Bites promote symptoms typical of viperid envenomation, such as pain, swelling and discoloration, all of which may be immediate. There are also reports of dizziness and tingling.^[3]

Humans respond rapidly to this venom, as do mice and birds. Lizards are less affected, while amphibians may even survive a bite. European snakes, such as Coronella and Natrix, are possibly immune.^[3]

V. ammodytes venom is used in the production of antivenin for the bite of other European vipers and the snake is farmed for this purpose.^[7][11]

Subspecies

Subspecies[5]	Taxon author[5]	Common name	Geographic range
V. a. ammodytes	(Linnaeus, 1758)	Western sand viper[10]	Austria (Styria, Carinthia), north Italy, Slovenia, Croatia, Bosnia-Herzegovina, Serbia, Montenegro, Macedonia, Albania, south-west Romania, north-west Bulgaria[3]
V. a. gregorwallneri	Sochurek, 1974		Austria, the former Yugoslavia[6]
V. a. meridionalis	Boulenger, 1903	Eastern sand viper[10]	Greece (incl. Corfu and other islands), Turkish Thrace[3]
V. a. montandoni	Boulenger, 1904	Transdanubian sand viper[10]	Bulgaria, south Romania[3]
V. a. transcaucasiana	Boulenger, 1913	Transcaucasian sand viper[3]	Georgia, north Turkish Anatolia[3]

Taxonomy

This species was originally described by Linnaeus in Systema Naturae in 1758. Subsequently, Boulenger described a number of subspecies in the early 20th century that are still mostly recognized today. However, there are many alternative taxonomies.^[3] One additional subspecies that may be encountered in literature is V. a. ruffoi (Bruno, 1968),^[3] found in the Alpine region of Italy. However, many consider both ruffoi and gregorwalineri to be synonymous with V. a. ammodytes^[6] and the taxon transcaucasiana to be a separate species.^[3][6]

See also

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

References

1. ^ ^{a b} 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} Street D. 1979. The Reptiles of Northern and Central Europe. London: B.T. Batsford Ltd. 268 pp. ISBN 0-7134-1374-3.
3. ^ ^{a b c d e f g h i j k l m n o p q r s t u v w x y z} Mallow D, Ludwig D, Nilson G. 2003. True Vipers: Natural History and Toxinology of Old World Vipers. Krieger Publishing Company. 359 pp. ISBN 0-89464-877-2.
4. ^ Gotch AF. 1986. Reptiles -- Their Latin Names Explained. Poole, UK: Blandford Press. 176 pp. ISBN 0-7137-1704-1.
5. ^ ^{a b c} "Vipera ammodytes". Integrated Taxonomic Information System. http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=634985. Retrieved 26 July 2006. 
6. ^ ^{a b c d e f g h} Strugariu A. 2006. The European Horn-Nosed Viper. VenomousReptiles.org.
7. ^ ^{a b} Mehrtens JM. 1987. Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
8. ^ U.S. Navy. 1991. Poisonous Snakes of the World. US Govt. New York: Dover Publications Inc. 203 pp. ISBN 0-486-26629-X.
9. ^ Convention on the Conservation of European Wildlife and Natural Habitats, Appendix II at Council of Europe. Accessed 9 October 2006.
10. ^ ^{a b c d} Steward JW. 1971. The Snakes of Europe. Cranbury, New Jersey: Associated University Press (Fairleigh Dickinson University Press). 238 pp. LCCCN 77-163307. ISBN 0-8386-1023-4.
11. ^ ^{a b c} Stidworthy J. 1974. Snakes of the world. Grosset & Dunlap Inc. ISBN 0-448-11856-4.
12. ^ Brown JH. 1973. Toxicology and Pharmacology of Venoms from Poisonous Snakes. Springfield, Illinois: Charles C. Thomas. 184 pp. LCCCN 73-229. ISBN 0-398-02808-7.

Further reading

• Arnold EN, Burton JA. 1978. A Field Guide to the Reptiles and Amphibians of Britain and Europe. London: Collins. 156 pp. ISBN 3-490-00318-7.
• Biella H-J. 1983. Die Sandotter. Die Neue Brehm-Bücherei. A Ziemsen Verlag. Wittenberg Lutherstadt. 84 pp.
• Bruno S. 1968. Sulla Vipera ammodytes in Italia. Memorie del Museo Civico di Storia Naturale, Verona, 15:289-386.
• Copley A, Banerjee S, Devi A. 1973. Studies of snake venom on blood coagulation. Part I: The thromboserpentin (thrombin-like) enzyme in the venoms. Thromb Res 2:487-508.
• Gulden J. 1988. Hibernation and breeding of V. ammodytes ammodytes. Litteratura Serpentium 8:168-72.
• Hays WST, Conant Sheila. 2007. Biology and Impacts of Pacific Island Invasive Species. 1. A Worldwide Review of Effects of the Small Indian Mongoose, Herpestes javanicus (Carnivora: Herpestidae) Pacific Science - Volume 61, Number 1, pp. 3–16.
• Nikolsky AM. 1916. Fauna of Russia and adjacent countries. Volume II: Ophidia. Petrograd. Translation from the Israel Program for Scientific Translations, Jerusalem, 1964, 247 pp.
• Mehrtens JM. 1987. Living Snakes of the World in Color. New York: Sterling Publishers. 256 pp. ISBN 0-8069-6460-X.
• Meier J, Stocker KF. 1991. Snake venom protein C activators. In: Tu A, editor. Reptile venoms and Toxins. New York: Marcel Dekker. pp 265–79.
• Mertens R, Wermuth H. 1960. Die Amphibien und Reptilien Europas. Verlag Waldemar Kramer, Frankfurt am Main, 1-264.
• McMahon M. 1990. Vipera ammodytes meridonalis envenomation. Journal of the Herpetological Association of Africa, 37:60.
• Petkovic D, Javanovic T, Micevic D, Unkovic-Cvetkovic N, Cvetkovic M. 1979. Action of Vipera ammodytes venom and its fractionation on the isolated rat heart. Toxicon, Great Britain, 17:639-44.
• Steward JW. 1971. The Snakes of Europe. London: David & Charles, Newton Abbot. 191 pp. ISBN 0-8386-1023-4.
• U.S. Navy. 1991. Poisonous Snakes of the World. New York: Dover Books. (Reprint of US Govt. Printing Office, Washington D.C.) 133 pp. ISBN 0-486-26629-X.

External links

• Vipera ammodytes at the Reptarium.cz Reptile Database. Accessed 21 November 2007.
• Vipera ammodytes at Amphibians and Reptiles of Europe. Accessed 7 October 2006.
• Vipera ammodytes - nose-horned viper at Faculty of Biology and Geology, "Babes-Bolyai" University, Romania. Accessed 7 October 2006.
• Vipera ammodytes at Club100. Accessed 7 October 2006.
• Vipera ammodytes^{[dead link]} at Amphibians & Reptiles in Bulgaria and Balkan Peninsula. Accessed 7 October 2006.
• Vipera ammodytes mating movie at VipersGarden. Accessed 7 October 2006.
• Video footage of three species, incl. V. ammodytes. on YouTube Accessed 4 November 2006.
• Vipera ammodytes giving birth in terrarium. pictures provided by Terrarium Morbidum. Accessed 4 June 2009.