Vipera ammodytes

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Viperoidea

Familia: Viperidae
Subfamilia: Viperinae
Genus: Vipera
Species: Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Viperoidea

Familia: Viperidae
Subfamilia: Viperinae
Genus: Vipera
Species: Vipera ammodytes
Subspecies (3): V. a. ammodytes – V. a. meridionalis – V. a. montandoni
Name

Vipera ammodytes (Linnaeus, 1758)

Holotype: UUZM 95.
Type locality: “Oriente”.

Combinations

Coluber ammodytes Linnaeus, 1758: 216 [original combination]
Vipera ammodytes — Sonnini & Latreille, 1820 [subsequent combination]

Synonyms

Vipera aspis balcanica Buresh & Zonkov, 1934

References
Primary references

Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio Decima, Reformata. Tomus I. Holmiæ (Stockholm): impensis direct. Laurentii Salvii. 824 pp. DOI: 10.5962/bhl.title.542 BHL Reference page.

Additional references

Schweiger, Mario 1992. Die Europäische Hornotter Vipera ammodytes (Linnaeus, 1758). Teil 1: Systematik, Ökologie und Lebensweise Herpetofauna 14 (77): 11–22

Links

Uetz, P. & Hallermann, J. 2022. Vipera ammodytes. The Reptile Database. Accessed on 7 June 2022.
Agasyan, A., Avci, A., Tuniyev, B, Isailovic, J.C., Lymberakis, P., Andrén, C., Cogalniceanu, D., Wilkinson, J., Ananjeva, N., Üzüm, N., Orlov, N., Podloucky, R., Tuniyev, S., Kaya, U., Sindaco, R., Böhme, W., Ajtic, R., Tok, V., Ugurtas, I.H., Sevinç, M., Tomović, L., Crochet, P.-A., Haxhiu, I., Joger, U., Sterijovski, B., Nilson, G. & Jelić, D. 2009. IUCN: Vipera ammodytes (Least Concern). The IUCN Red List of Threatened Species 2009: e.T62255A12584303. DOI: 10.2305/IUCN.UK.2009.RLTS.T62255A12584303.en. Accessed on 07 June 2022.

Vernacular names
български: Пепелянка
brezhoneg: Naer-wiber gornek
bosanski: Poskok
čeština: Zmije růžkatá
Deutsch: Europäische Hornotter
Ελληνικά: οχιά
English: Nose-horned Viper
español: Víbora cornuda
eesti: Nokisrästik
euskara: Sugegorri sudurluze
suomi: Hietakyy
français: vipère cornue
hrvatski: Poskok
magyar: Homoki vipera
italiano: Vipera dal corno
日本語: ハナダカクサリヘビ
ქართული: ცხვირრქოსანი გველგესლა
lietuvių: Ilganosė angis
македонски: Поскок
Nederlands: Zandadder
polski: Żmija nosoroga
русский: Носатая гадюка
slovenščina: Modras
српски / srpski: Поскок
svenska: Sandhuggorm
Türkçe: Boynuzlu engerek
українська: Гадюка носата

Vipera ammodytes (other common names include horned viper, long-nosed viper, nose-horned viper, sand viper)[3] is a venomous viper species found in southern Europe, mainly northern Italy, 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.[4] The specific name, ammodytes, is derived from the Greek words ammos, meaning "sand", and dutes, meaning "burrower" or "diver", despite its preference for rocky habitats.[5] Five subspecies are currently recognized, including the nominate subspecies described here.[6]

Description

V. ammodytes grows to a maximum total length (body + tail) of 95 cm (37.5 in), although individuals usually measure less than 85 cm (33.5 in). Maximum length also depends on locality,[vague] with northern forms distinctly larger than southern ones.[4] The average total length is 50–70 cm (19.5–27.5 in) with reports of specimens over 1 m (39 in) in total length.[7] Females are usually[vague] larger and more heavily[specify] built, although the largest specimens on record are males.[7]

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 scales 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.[4]

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.[4] It grows to a length of about 5 mm (0.20 in) and is actually soft and flexible. In southern subspecies, the horn sits vertically upright, while in V. a. Ammodytes it points diagonally forward.[3]

The body is covered with strongly keeled dorsal scales in 21 or 23 rows (rarely 25) at 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.[4]
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 gray, sometimes yellowish or pinkish gray, or yellowish brown. The dorsal zigzag is dark gray 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.[4]

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.[4]

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.[4] The cinderella snake may be between the rail and a ballast prism, accumulating solar heat in her body .
Common names

The common names that can be found for Vipera ammodytes in English are horned viper, long-nosed viper, nose-horned viper, sand viper,[3] sand adder, common sand adder, common sand viper,[8] and sand natter.[9] In Croatia, Bosnia and Herzegovina, North Macedonia, and Serbia, a common name used for this snake is poskok (поскок).
Geographic range

North-eastern Italy, Slovenia, Croatia, Bosnia-Herzegovina, Serbia, Montenegro, Albania, Kosovo, North Macedonia, Greece (including Cyclades), southern Austria, Romania, Bulgaria, Turkey, Georgia and Syria.
Vipera ammodytes distribution (including V. a. transcaucasiana)

The type locality is listed as "Oriente". Schwarz (1936) proposed that the type locality be restricted to "Zara" (Zadar, Croatia).[2]
Conservation status

This species is listed as strictly protected (Appendix II) under the Berne Convention.[10]
Habitat

The common name sand viper is misleading, as this species does not occur in mostly sandy areas.[11] V. ammodytes primarily 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.[4]
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.[4]

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.[4]

Vipera ammodytes hibernates in the winter for a period of 2 to 6 months depending on environmental conditions.[7]
V. ammodytes
Feeding

Adults generally feed on small mammals and birds, whilst juveniles predominantly eat lizards. Feeding behavior changes and is influenced heavily 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.[4] There are also reports of cannibalism.[7] Arthropods such as large insects and centipedes have regularly been found in the stomach contents of vipers, albeit more frequently in juveniles that eat centipedes like Mediterranean banded centipede.[12]
Reproduction

Before mating, the males of this species will engage in a combat dance, similar to adders.[4] Mating takes place in the spring (April–May), and one to twenty live young are born in late summer or fall (August–October). At birth, juveniles are 14–24 cm (5.5–9.4 in) in total length.[7] This species is ovoviviparous.[13]
Captivity

This species has often been kept in captivity and bred successfully.[4] It tolerates captivity much better than other European vipers, thriving in most surroundings and usually taking food easily from the start.[13] 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.[7]
Venom

This is likely the most dangerous snake to be found in Europe.[14] In some areas it is at least a significant medical risk.

The venom can be quite toxic [based on tests conducted solely on mice], but varies over time and among different populations.[4] Brown (1973) gives an LD50 for mice of 1.2 mg/kg IV, 1.5 mg/kg IP and 2.0 mg/kg SC.[15] 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.[4]

The venom has both proteolytic and neurotoxic components and contains hemotoxins with blood coagulant properties, similar to and as powerful as in crotaline 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.[4] The venom is rarely deadly when hospital care is available.[16]

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.[4]

Vipera ammodytes venom is used in the production of antivenin for the bite of other European vipers and the snake is farmed for this purpose.[8][13]
Taxonomy

This species was originally described by Carl Linnaeus in Systema Naturae in 1758. Subsequently, George Albert Boulenger described a number of subspecies in the early 20th century that are still mostly recognized today. However, there are many alternative taxonomies.[4] One additional subspecies that may be encountered in literature is V. a. ruffoi (Bruno, 1968),[4] found in the Alpine region of Italy. However, many consider both V. a. ruffoi and V. a. gregorwalineri to be synonymous with V. a. ammodytes,[7] and consider V. a. transcaucasiana to be a separate species.[4][7]
Subspecies[6] Taxon author[6] Common name Geographic range
V. a. ammodytes (Linnaeus, 1758) Western sand viper[11] Austria (Styria, Carinthia), north Italy, Slovenia, Croatia, Bosnia-Herzegovina, Serbia, Montenegro, North Macedonia, Albania, south-west Romania, north-west Bulgaria[4]
V. a. gregorwallneri Sochurek, 1974 Austria, Slovenia, Croatia, Bosnia-Herzegovina, Serbia, Montenegro, North Macedonia[7]
V. a. meridionalis Boulenger, 1903 Eastern sand viper[11] Greece (incl. Corfu and other islands), Turkish Thrace[4]
V. a. montandoni Boulenger, 1904 Transdanubian sand viper[11] Bulgaria, south Romania[4]
V. a. transcaucasiana Boulenger, 1913 Transcaucasian sand viper[4] Georgia, north Turkish Anatolia[4]
References

Aram Agasyan; Aziz Avci; Boris Tuniyev; Jelka Crnobrnja Isailovic; Petros Lymberakis; Claes Andrén; Dan Cogalniceanu; John Wilkinson; Natalia Ananjeva; Nazan Üzüm; et al. (2009). "Vipera ammodytes". IUCN Red List of Threatened Species. 2009: e.T62255A12584303. doi:10.2305/IUCN.UK.2009.RLTS.T62255A12584303.en. Retrieved 12 June 2022.
McDiarmid, R.W., Campbell, J.A., Touré, T. 1999. Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
Street, D. 1979. The Reptiles of Northern and Central Europe. London: B.T. Batsford Ltd. 268 pp. ISBN 0-7134-1374-3.
Mallow, D., Ludwig, D., Nilson, G. 2003. True Vipers: Natural History and Toxinology of Old World Vipers. Malabar, Florida: Krieger Publishing Company. 359 pp. ISBN 0-89464-877-2.
Gotch, A.F. 1986. Reptiles – Their Latin Names Explained. Poole, UK: Blandford Press. 176 pp. ISBN 0-7137-1704-1.
"Vipera ammodytes (Linnaeus, 1758)". Integrated Taxonomic Information System. Retrieved 26 July 2006.
Strugariu, A. 2006. The European Horn-Nosed Viper Archived 14 October 2013 at the Wayback Machine. VenomousReptiles.org.
Mehrtens, J.M. 1987. Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
U.S. Navy. 1991. Poisonous Snakes of the World. US Govt. New York: Dover Publications Inc. 203 pp. ISBN 0-486-26629-X.
Convention on the Conservation of European Wildlife and Natural Habitats, Appendix II Archived 16 July 2015 at the Wayback Machine at Council of Europe. Accessed 9 October 2006.
Steward, J.W. 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.
Plettenberg-Laing, Anthony (2020). "Observations on the diet of the nose-horned viper (Vipera ammodytes) in Greece". Herpetological Bulletin. 153 (153, Autumn 2020): 37–39. doi:10.33256/hb153.3739.
Stidworthy, J. 1974. Snakes of the World. Grosset & Dunlap Inc. ISBN 0-448-11856-4.
Arnold, E.N.; Burton, J.A. 1978. A Field Guide to the Reptiles and Amphibians of Britain and Europe. London: Collins. 272 pp. ISBN 0-00-219318-3. (Vipera ammodytes, pp. 222–223 + Plate 40 + Map 125.)
Brown, J.H. 1973. Toxicology and Pharmacology of Venoms from Poisonous Snakes. Springfield, Illinois: Charles C. Thomas. 184 pp. LCCCN 73–229. ISBN 0-398-02808-7.

Radonić, V.; Budimir, D.; Bradarić, N.; Luksić, B.; Sapunar, D.; Vilović, K. (1997). "Envenomation by the Horned Viper (Vipera ammodytes L.)" (PDF). Military Medicine. 162 (3): 179–182. doi:10.1093/milmed/162.3.179. PMID 9121663. Retrieved 29 July 2022.