Vipera ammodytes
Encyclopedia
Common names: horned viper, long-nosed viper, nose-horned viper, sand viper, more.

Vipera ammodytes is a venomous viper
Viperinae
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...

 species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 found in southern Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

 through to the Balkans
Balkans
The Balkans is a geopolitical and cultural region of southeastern Europe...

 and parts of the Middle East
Middle East
The Middle East is a region that encompasses Western Asia and Northern Africa. It is often used as a synonym for Near East, in opposition to Far 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. The specific name is derived from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 words ammos and dutes, meaning "sand" and "burrower" or "diver"; not a very good name for an animal that actually prefers rocky habitats. Five subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

 are currently recognized, including the nominate subspecies described here.

Description

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. 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.

The head is covered in small, irregular scales that are either smooth or only weakly keeled, except for a pair of large supraocular scales
Supraocular scales
In scaled reptiles, supraocular scales are scales on the crown immediately above the eye. The size and shape of these scales are among the many characteristics used to differentiate species from each another....

 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
Nasorostral scale
In reptiles, the nasorostral is an enlarged and usually paired scale, just behind the rostral ....

. The rostral scale is wider than it is long.

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. 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.

The body is covered with strongly keeled
Keeled scales
Keeled scales refer to reptile scales that, rather than being smooth, have a ridge down the center that may or may not extend to the tip of the scale, making them rough to the touch...

 dorsal scales
Dorsal scales
In snakes, the dorsal scales are the longitudinal series of plates that encircle the body, but do not include the ventral scales.When counting dorsal scales, numbers are often given for three points along the body, for example 19:21:17...

 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
Anal scale
In snakes, the anal scale is the scale just in front of and covering the cloacal opening. This scale can be either single or paired . When paired, the division is oblique. It is preceded by the ventral scales and followed by the subcaudal scales....

 is single.

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.

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.

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.

Common names

Horned viper, long-nosed viper, nose-horned viper, sand viper, sand adder, common sand adder, common sand viper, sand natter.

Geographic range

Southern Austria
Austria
Austria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the...

, north-eastern Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

, Slovenia
Slovenia
Slovenia , officially the Republic of Slovenia , is a country in Central and Southeastern Europe touching the Alps and bordering the Mediterranean. Slovenia borders Italy to the west, Croatia to the south and east, Hungary to the northeast, and Austria to the north, and also has a small portion of...

, Croatia
Croatia
Croatia , officially the Republic of Croatia , is a unitary democratic parliamentary republic in Europe at the crossroads of the Mitteleuropa, the Balkans, and the Mediterranean. Its capital and largest city is Zagreb. The country is divided into 20 counties and the city of Zagreb. Croatia covers ...

, Bosnia-Herzegovina, Serbia
Serbia
Serbia , officially the Republic of Serbia , is a landlocked country located at the crossroads of Central and Southeast Europe, covering the southern part of the Carpathian basin and the central part of the Balkans...

, Montenegro
Montenegro
Montenegro Montenegrin: Crna Gora Црна Гора , meaning "Black Mountain") is a country located in Southeastern Europe. It has a coast on the Adriatic Sea to the south-west and is bordered by Croatia to the west, Bosnia and Herzegovina to the northwest, Serbia to the northeast and Albania to the...

, Albania
Albania
Albania , officially known as the Republic of Albania , is a country in Southeastern Europe, in the Balkans region. It is bordered by Montenegro to the northwest, Kosovo to the northeast, the Republic of Macedonia to the east and Greece to the south and southeast. It has a coast on the Adriatic Sea...

, Republic of Macedonia
Republic of Macedonia
Macedonia , officially the Republic of Macedonia , is a country located in the central Balkan peninsula in Southeast Europe. It is one of the successor states of the former Yugoslavia, from which it declared independence in 1991...

, Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....

 (including Greek Macedonia
Macedonia (Greece)
Macedonia is a geographical and historical region of Greece in Southern Europe. Macedonia is the largest and second most populous Greek region...

 and Cyclades
Cyclades
The Cyclades is a Greek island group in the Aegean Sea, south-east of the mainland of Greece; and a former administrative prefecture of Greece. They are one of the island groups which constitute the Aegean archipelago. The name refers to the islands around the sacred island of Delos...

), Romania
Romania
Romania is a country located at the crossroads of Central and Southeastern Europe, on the Lower Danube, within and outside the Carpathian arch, bordering on the Black Sea...

, Bulgaria
Bulgaria
Bulgaria , officially the Republic of Bulgaria , is a parliamentary democracy within a unitary constitutional republic in Southeast Europe. The country borders Romania to the north, Serbia and Macedonia to the west, Greece and Turkey to the south, as well as the Black Sea to the east...

, Turkey
Turkey
Turkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...

, Georgia
Georgia (country)
Georgia is a sovereign state in the Caucasus region of Eurasia. Located at the crossroads of Western Asia and Eastern Europe, it is bounded to the west by the Black Sea, to the north by Russia, to the southwest by Turkey, to the south by Armenia, and to the southeast by Azerbaijan. The capital of...

 and Syria
Syria
Syria , officially the Syrian Arab Republic , is a country in Western Asia, bordering Lebanon and the Mediterranean Sea to the West, Turkey to the north, Iraq to the east, Jordan to the south, and Israel to the southwest....

. The type locality is listed as "Oriente." Schwartz (1936) proposed that the type locality be restricted to "Zara" (Zadar
Zadar
Zadar is a city in Croatia on the Adriatic Sea. It is the centre of Zadar county and the wider northern Dalmatian region. Population of the city is 75,082 citizens...

, Croatia).

Conservation status

This species is listed as strictly protected (Appendix II) under the Berne Convention
Convention on the Conservation of European Wildlife and Natural Habitats
The Bern Convention on the Conservation of European Wildlife and Natural Habitats 1979, also known as the Bern Convention , came into force on June 1, 1982....

.

Habitat

The common name sand viper is misleading, as this species does not occur in really sandy areas. 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.

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.

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.

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

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. There are also reports of cannibalism.

Reproduction

Before mating, the males of this species will engage an a combat dance, similar to adders. 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. This species is ovoviviparous
Ovoviviparity
Ovoviviparity, ovovivipary, or ovivipary, is a mode of reproduction in animals in which embryos develop inside eggs that are retained within the mother's body until they are ready to hatch...

.

Captivity

This species has often been kept in captivity and bred successfully. It tolerates captivity much better than other European vipers, thriving in most surroundings and usually takes food easily from the start. 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.

Venom

This is likely the most dangerous snake to be found in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

. In some areas it is at least a significant medical risk; in the past fatalities were relatively frequent in the Balkans
Balkans
The Balkans is a geopolitical and cultural region of southeastern Europe...

 because the peasants there had a habit of walking barefoot.

The venom can be quite toxic
Toxicity
Toxicity is the degree to which a substance can damage a living or non-living organisms. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a substructure of the organism, such as a cell or an organ , such as the liver...

 [based on tests conducted solely on mice], but varies over time and among different populations. Brown (1973) gives an for mice of 1.2 mg/kg IV
Intravenous therapy
Intravenous therapy or IV therapy is the infusion of liquid substances directly into a vein. The word intravenous simply means "within a vein". Therapies administered intravenously are often called specialty pharmaceuticals...

, 1.5 mg/kg IP and 2.0 mg/kg SC. 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.

The venom has both proteolytic
Proteolysis
Proteolysis is the directed degradation of proteins by cellular enzymes called proteases or by intramolecular digestion.-Purposes:Proteolysis is used by the cell for several purposes...

 and neurotoxic
Neurotoxin
A neurotoxin is a toxin that acts specifically on nerve cells , usually by interacting with membrane proteins such as ion channels. Some sources are more general, and define the effect of neurotoxins as occurring at nerve tissue...

 components and contains hemotoxin
Hemotoxin
Hemotoxins, haemotoxins or hematotoxins are toxins that destroy red blood cells , disrupt blood clotting, and/or cause organ degeneration and generalized tissue damage. The term hemotoxin is to some degree a misnomer since toxins that damage the blood also damage other tissues...

s with blood coagulant properties, similar to and as powerful as in crotalid
Crotalinae
The Crotalinae, commonly known as "pit vipers" or crotaline snakes, are a subfamily of venomous vipers found in Asia and the Americas. They are distinguished by the presence of a heat-sensing pit organ located between the eye and the nostril on either side of the head...

 venom. Other properties include anticoagulant effects, hemoconcentration and hemorrhage
Bleeding
Bleeding, technically known as hemorrhaging or haemorrhaging is the loss of blood or blood escape from the circulatory system...

. Bites promote symptoms typical of viperid
Viperidae
The Viperidae are a family of venomous snakes found all over the world, except in Antarctica, Australia, Ireland, Madagascar, Hawaii, various other isolated islands, and above the Arctic Circle. All have relatively long, hinged fangs that permit deep penetration and injection of venom. Four...

 envenomation, such as pain, swelling and discoloration, all of which may be immediate. There are also reports of dizziness and tingling.

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
Coronella
Coronella is a genus of harmless colubrids found in Europe, North Africa and West Asia. Three species are currently recognized.-Description:...

 and Natrix
Natrix
Natrix is a genus of colubrid snakes. There are three or four species in the genus. They are collectively called grass snakes and water snakes...

, are possibly immune.

V. ammodytes venom is used in the production of antivenin
Antivenin
Antivenom is a biological product used in the treatment of venomous bites or stings. Antivenom is created by milking venom from the desired snake, spider or insect. The venom is then diluted and injected into a horse, sheep or goat...

 for the bite of other European vipers and the snake is farmed for this purpose.

Subspecies

Subspecies Taxon author Common name Geographic range
V. a. ammodytes (Linnaeus
Carolus Linnaeus
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...

, 1758)
Western sand viper Austria
Austria
Austria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the...

 (Styria, Carinthia
Carinthia (state)
Carinthia is the southernmost Austrian state or Land. Situated within the Eastern Alps it is chiefly noted for its mountains and lakes.The main language is German. Its regional dialects belong to the Southern Austro-Bavarian group...

), north Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

, Slovenia
Slovenia
Slovenia , officially the Republic of Slovenia , is a country in Central and Southeastern Europe touching the Alps and bordering the Mediterranean. Slovenia borders Italy to the west, Croatia to the south and east, Hungary to the northeast, and Austria to the north, and also has a small portion of...

, Croatia
Croatia
Croatia , officially the Republic of Croatia , is a unitary democratic parliamentary republic in Europe at the crossroads of the Mitteleuropa, the Balkans, and the Mediterranean. Its capital and largest city is Zagreb. The country is divided into 20 counties and the city of Zagreb. Croatia covers ...

, Bosnia-Herzegovina, Serbia
Serbia
Serbia , officially the Republic of Serbia , is a landlocked country located at the crossroads of Central and Southeast Europe, covering the southern part of the Carpathian basin and the central part of the Balkans...

, Montenegro
Montenegro
Montenegro Montenegrin: Crna Gora Црна Гора , meaning "Black Mountain") is a country located in Southeastern Europe. It has a coast on the Adriatic Sea to the south-west and is bordered by Croatia to the west, Bosnia and Herzegovina to the northwest, Serbia to the northeast and Albania to the...

, Macedonia
Republic of Macedonia
Macedonia , officially the Republic of Macedonia , is a country located in the central Balkan peninsula in Southeast Europe. It is one of the successor states of the former Yugoslavia, from which it declared independence in 1991...

, Albania
Albania
Albania , officially known as the Republic of Albania , is a country in Southeastern Europe, in the Balkans region. It is bordered by Montenegro to the northwest, Kosovo to the northeast, the Republic of Macedonia to the east and Greece to the south and southeast. It has a coast on the Adriatic Sea...

, south-west Romania
Romania
Romania is a country located at the crossroads of Central and Southeastern Europe, on the Lower Danube, within and outside the Carpathian arch, bordering on the Black Sea...

, north-west Bulgaria
Bulgaria
Bulgaria , officially the Republic of Bulgaria , is a parliamentary democracy within a unitary constitutional republic in Southeast Europe. The country borders Romania to the north, Serbia and Macedonia to the west, Greece and Turkey to the south, as well as the Black Sea to the east...

V. a. gregorwallneri
Vipera ammodytes gregorwallneri
Vipera ammodytes gregorwallneri is a of venomous viper subspecies found mainly in Austria and the former Yugoslavia.-Taxonomy:Many authors, such as Golay et al. do not recognize this taxon and instead relegate it to the synonymy of V. a. ammodytes....

Sochurek, 1974 Austria, the former Yugoslavia
Yugoslavia
Yugoslavia refers to three political entities that existed successively on the western part of the Balkans during most of the 20th century....

V. a. meridionalis
Vipera ammodytes meridionalis
Vipera ammodytes meridionalis is a venomous viper subspecies found in Greece and Turkish Thrace.-Description:According to Boulenger : "Naso-rostral shield never reaching the canthus rostralis and but rarely extending higher up than the upper border of the rostral, which is often as deep as broad,...

Boulenger
George Albert Boulenger
George Albert Boulenger FRS was a Belgian-British zoologist who identified over 2000 new animal species, chiefly fish, reptiles and amphibians.-Life:...

, 1903
Eastern sand viper Greece (incl. Corfu
Corfu
Corfu is a Greek island in the Ionian Sea. It is the second largest of the Ionian Islands, and, including its small satellite islands, forms the edge of the northwestern frontier of Greece. The island is part of the Corfu regional unit, and is administered as a single municipality. The...

 and other islands), Turkish Thrace
Thrace
Thrace is a historical and geographic area in southeast Europe. As a geographical concept, Thrace designates a region bounded by the Balkan Mountains on the north, Rhodope Mountains and the Aegean Sea on the south, and by the Black Sea and the Sea of Marmara on the east...

V. a. montandoni
Vipera ammodytes montandoni
Vipera ammodytes montandoni is a venomous viper subspecies found in Bulgaria and southern Romania.-Description:According to Boulenger : "Naso-rostral shield never reaching the canthus rostralis nor the summit of the rostral shield, which is deeper than broad ; rostral appendage clad with ten to...

Boulenger, 1904 Transdanubian sand viper Bulgaria, south Romania
V. a. transcaucasiana
Vipera ammodytes transcaucasiana
Vipera ammodytes transcaucasiana is a venomous viper subspecies found in parts of Georgia and northern Turkish Anatolia.-Description:Grows to a maximum length of 75 cm, but is usually not so large....

Boulenger, 1913 Transcaucasian sand viper Georgia, north Turkish Anatolia

Taxonomy

This species was originally described by Linnaeus
Carolus Linnaeus
Carl Linnaeus , also known after his ennoblement as , was a Swedish botanist, physician, and zoologist, who laid the foundations for the modern scheme of binomial nomenclature. He is known as the father of modern taxonomy, and is also considered one of the fathers of modern ecology...

 in Systema Naturae
Systema Naturae
The book was one of the major works of the Swedish botanist, zoologist and physician Carolus Linnaeus. The first edition was published in 1735...

 in 1758. Subsequently, Boulenger
George Albert Boulenger
George Albert Boulenger FRS was a Belgian-British zoologist who identified over 2000 new animal species, chiefly fish, reptiles and amphibians.-Life:...

described a number of subspecies in the early 20th century that are still mostly recognized today. However, there are many alternative taxonomies. One additional subspecies that may be encountered in literature is V. a. ruffoi (Bruno, 1968), found in the Alpine region of Italy. However, many consider both ruffoi and gregorwalineri to be synonymous with V. a. ammodytes and the taxon transcaucasiana to be a separate species.

External links

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