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The puff adder (''Bitis arietans'') is a highly venomous
viper Vipers are snakes in the family Viperidae, found in most parts of the world, except for Antarctica, Australia, Hawaii, Madagascar, New Zealand, Ireland, and various other isolated islands. They are venomous and have long (relative to non-vipe ...
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
found in
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
s and
grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...
s from
Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
and western
Arabia The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
throughout
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
except for the
Sahara The Sahara (, ) is a desert spanning across North Africa. With an area of , it is the largest hot desert in the world and the list of deserts by area, third-largest desert overall, smaller only than the deserts of Antarctica and the northern Ar ...
and
rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...
regions. It is responsible for causing the most snakebite fatalities in Africa owing to various factors, such as its wide distribution, frequent occurrence in highly populated regions, and aggressive disposition.. Like all other vipers, it is
venomous Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
. Two
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
are currently recognized, including the
nominate subspecies In biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. ...
described here. The species is commonly known as the puff adder,. African puff adder,. or common puff adder.


Taxonomy

German naturalist
Blasius Merrem Blasius Merrem (4 February 1761 – 23 February 1824) was a German natural history, naturalist, zoologist, Ornithology, ornithologist, mathematician, and Herpetology, herpetologist. In 1804, he became the professor of political economy and botan ...
described the puff adder in 1820. The word ''arietans'' means "striking violently" and is derived from the Latin ''arieto''. The type locality given is "''Promontorio bonae spei''" (
Cape of Good Hope The Cape of Good Hope ( ) is a rocky headland on the Atlantic Ocean, Atlantic coast of the Cape Peninsula in South Africa. A List of common misconceptions#Geography, common misconception is that the Cape of Good Hope is the southern tip of Afri ...
),
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
.


Subspecies


Description

The snake's typical size is about 1.0 m (39.3 in) in total length (body and tail) and very stout. Large specimens of 190 cm (75 in) total length, weighing over 6.0 kg (13.2 lb) and with a girth of 40 cm (16 in) have been reported. Specimens from Saudi Arabia are not as large, usually no more than in total length. Males are usually larger than females and have relatively longer tails. The color pattern varies geographically. The head has two well-marked dark bands – one on the crown and the other between the eyes. On the sides of the head, two oblique, dark bands or bars run from the eye to the supralabials. Below, the head is yellowish white with scattered dark blotches. Iris color ranges from gold to silver-gray. Dorsally, the ground color varies from straw yellow, to light brown, to orange or reddish brown. This is overlaid with a pattern of 18–22 backwardly directed, dark brown to black bands that extend down the back and tail. Usually, these bands are roughly chevron-shaped, but may be more U-shaped in some areas. They also form two to six light-and-dark cross-bands on the tail. Some populations are heavily flecked with brown and black, often obscuring other coloration, giving the animal a dusty-brown or blackish appearance. The belly is yellow or white, with a few scattered dark spots. Newborn young have golden head markings with pinkish to reddish ventral plates toward the lateral edges.
One unusual specimen, described by Branch and Farrell (1988), from Summer Pride,
East London East London is the part of London, England, east of the ancient City of London and north of the River Thames as it begins to widen. East London developed as London Docklands, London's docklands and the primary industrial centre. The expansion of ...
, in South Africa, was striped. The pattern consisted of a narrow (one scale wide), pale yellowish stripe that ran from the crown of the head to the tip of the tail.
Generally, though, these are relatively dull-looking snakes, except for male specimens from highland East Africa and the
Western Cape The Western Cape ( ; , ) is a provinces of South Africa, province of South Africa, situated on the south-western coast of the country. It is the List of South African provinces by area, fourth largest of the nine provinces with an area of , an ...
province of South Africa, that usually have a striking yellow-and-black color pattern. Puff adders have a form of olfactory crypsis, which has been shown to make detecting them difficult for trained dogs and meerkats, both scent-based predators. The exact nature of this ability is not known, but is hypothesized to be related to a low metabolic rate, as well as relocation after shedding and defecating.


Scalation

The head has a less than triangular shape with a blunt and rounded snout. Still, the head is much wider than the neck. The
rostral scale The rostral scale, or rostral, in snakes and other scaled reptiles is the median plate on the tip of the snout that borders the mouth opening. Wright AH, Wright AA (1957). ''Handbook of Snakes of the United States and Canada''. Ithaca and London: ...
is small. The circumorbital ring consists of 10–16 scales. Across the top of the head, there are 7–11 interocular scales; three or four scales separate the suboculars and the
supralabials In reptiles, the supralabial scales, also called upper-labials, are those scales that border the mouth opening along the upper jaw. They do not include the median scaleWright AH, Wright AA. 1957. Handbook of Snakes. Comstock Publishing Associates ( ...
. It has 12 to 17
supralabials In reptiles, the supralabial scales, also called upper-labials, are those scales that border the mouth opening along the upper jaw. They do not include the median scaleWright AH, Wright AA. 1957. Handbook of Snakes. Comstock Publishing Associates ( ...
and 13–17 sublabials. The first three or four sublabials contact the chin shields, of which only one pair exists. Often, two fangs are on each
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
, and both can be functional. Midbody, the snake has 29–41 rows of
dorsal scales In snakes, the dorsal scales are the longitudinal series of plates that encircle the body, but do not include the ventral scales. Campbell JA, Lamar WW (2004). ''The Venomous Reptiles of the Western Hemisphere''. Ithaca and London: Comstock Publis ...
. These are strongly keeled except for the outermost rows. The ventral scale count is 123–147, the subcaudals number 14–38. Females have no more than 24 subcaudals. The anal scale is single.


Distribution and habitat

This species may be the most common and widespread snake in Africa. It is found in most African regions and on parts of the
Arabian Peninsula The Arabian Peninsula (, , or , , ) or Arabia, is a peninsula in West Asia, situated north-east of Africa on the Arabian plate. At , comparable in size to India, the Arabian Peninsula is the largest peninsula in the world. Geographically, the ...
. It is found in all habitats except true deserts, rainforests, and (tropical) alpine habitats. It is most often associated with rocky grasslands.Mehrtens JM. 1987. ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. . It is not found in rainforest areas, such as along the coast of
West Africa West Africa, also known as Western Africa, is the westernmost region of Africa. The United Nations geoscheme for Africa#Western Africa, United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Gha ...
and in
Central Africa Central Africa (French language, French: ''Afrique centrale''; Spanish language, Spanish: ''África central''; Portuguese language, Portuguese: ''África Central'') is a subregion of the African continent comprising various countries accordin ...
(i.e., central DR Congo); it is also absent from the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
coastal region of
North Africa North Africa (sometimes Northern Africa) is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region. However, it is sometimes defined as stretching from the Atlantic shores of t ...
. On the Arabian Peninsula, it is found as far north as
Ta'if Taif (, ) is a city and governorate in Mecca Province in Saudi Arabia. Located at an elevation of in the slopes of the Hijaz Mountains, which themselves are part of the Sarawat Mountains, Sarat Mountains, the city has a population of 563,282 pe ...
. It has been reported to be found in the Dhofar region of southern
Oman Oman, officially the Sultanate of Oman, is a country located on the southeastern coast of the Arabian Peninsula in West Asia and the Middle East. It shares land borders with Saudi Arabia, the United Arab Emirates, and Yemen. Oman’s coastline ...
.


Behaviour

Normally a sluggish species, the puff adder relies on
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
for protection. Locomotion is primarily rectilinear, using the broad ventral scales in a caterpillar fashion and aided by its own weight for traction. When agitated, it can resort to a typical serpentine movement of surprising speed. Although mainly terrestrial, these snakes are good swimmers and can also climb with ease; often they are found basking in low bushes. One specimen was found 4.6 m above the ground in a densely branched tree. If disturbed, they hiss loudly and continuously, adopting a tightly coiled defensive posture with the forepart of their body held in a taut "S" shape. At the same time, they may attempt to back away from the threat towards cover. They may strike suddenly and fast, to the side as easily as forwards, before returning quickly to the defensive position, ready to strike again. During a strike, the force of the impact is so strong, and the long fangs penetrate so deeply, that prey items are often killed by the physical trauma alone. The fangs apparently can penetrate soft leather. They can strike to a distance of about one-third of their body length, but juveniles can launch their entire bodies forwards in the process. These snakes rarely grip their victims, but instead release quickly to return to the striking position.


Feeding

Mostly nocturnal, they rarely forage actively, preferring, instead, to ambush prey as it happens by. Their prey includes mammals (rodents 'Aethomys'' sp., ''Arvicanthis">Aethomys.html" ;"title="'Aethomys">'Aethomys'' sp., ''Arvicanthis'' sp., ''Mastomys'' sp., ''Otomys'' sp., ''Rattus'' sp., ''Rhabdomys pumilio'', and ''Saccostomus campestris''] and even occasionally small deer), birds, amphibians (such as ''Schismaderma carens''), lizards, other snakes, and tortoises.


Reproduction

Females produce a pheromone to attract males, which engage in neck-wrestling combat dances. A female in
Malindi Malindi is a town on Malindi Bay at the mouth of the Sabaki River, lying on the Indian Ocean coast of Kenya. It is 120 kilometres northeast of Mombasa. The population of Malindi was 119,859 as of the 2019 census. It is the largest urban centr ...
was followed by seven males. They give birth to large numbers of offspring; litters over 80 have been reported, while 50–60 are not unusual. Newborns are in length. Very large specimens, particularly those from East Africa, give birth to the highest numbers of offspring. A Kenyan female in a
Czech Czech may refer to: * Anything from or related to the Czech Republic, a country in Europe ** Czech language ** Czechs, the people of the area ** Czech culture ** Czech cuisine * One of three mythical brothers, Lech, Czech, and Rus *Czech (surnam ...
zoo gave birth to 156 young, the largest litter for any species of snake.


Captivity

These snakes do well in captivity, but gluttony has been reported. Kauffeld (1969) mentions that specimens can be maintained for years on only one meal per week, but that when offered all they can eat, the result is often death, or at best wholesale regurgitation. They are bad-tempered snakes, and some specimens never settle down in captivity, always hissing and puffing when approached.


Venom

This species is responsible for more snakebite fatalities than any other African snake, due to a combination of factors, including its wide distribution, common occurrence, large size, potent venom that is produced in large amounts, long fangs, and habit of basking by footpaths and sitting quietly when approached. The venom has
cytotoxic Cytotoxicity is the quality of being toxic to cells. Examples of toxic agents are toxic metals, toxic chemicals, microbe neurotoxins, radiation particles and even specific neurotransmitters when the system is out of balance. Also some types of dr ...
effects and is one of the most
toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subst ...
of any vipers based on LD50. The values in mice vary: 0.4–2.0 mg/kg
intravenous Intravenous therapy (abbreviated as IV therapy) is a medical technique that administers fluids, medications and nutrients directly into a person's vein. The intravenous route of administration is commonly used for rehydration or to provide nutr ...
ly, 0.9–3.7 mg/kg peritoneally, and 4.4–7.7 mg/kg subcutaneously (SC). Mallow et al. (2003) give an LD50 range of 1.0–7.75 mg/kg SC. Venom yield is typically 150–350 mg, with a maximum of 750 mg. Brown (1973) mentions a venom yield of 180–750 mg. About are thought to be enough to kill a healthy adult human male, with death occurring after 25 hours. In humans, bites from this species can produce severe local and systemic symptoms. Based on the degree and type of local effect, bites can be divided into two symptomatic categories – those with little or no surface extravasation, and those with
hemorrhage Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, ...
s evident as ecchymosis, bleeding, and swelling. In both cases, severe pain and tenderness occur, but in the latter, widespread superficial or deep
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
and
compartment syndrome Compartment syndrome is a serious medical condition in which increased pressure within a Fascial compartment, body compartment compromises blood flow and tissue function, potentially leading to permanent damage if not promptly treated. There are ...
are seen. Serious bites cause limbs to become immovably flexed as a result of significant
hemorrhage Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, ...
or
coagulation Coagulation, also known as clotting, is the process by which blood changes from a liquid to a gel, forming a thrombus, blood clot. It results in hemostasis, the cessation of blood loss from a damaged vessel, followed by repair. The process of co ...
in the affected muscles. Residual induration, however, is rare and usually these areas completely resolve. Other bite symptoms that may occur in humans include
edema Edema (American English), also spelled oedema (British English), and also known as fluid retention, swelling, dropsy and hydropsy, is the build-up of fluid in the body's tissue (biology), tissue. Most commonly, the legs or arms are affected. S ...
, which may become extensive, shock, watery blood oozing from the puncture wounds, nausea and vomiting, subcutaneous bruising, blood blisters that may form rapidly, and painful swelling of the regional lymph nodes. Swelling usually decreases after a few days, except for the area immediately around the bite site.
Hypotension Hypotension, also known as low blood pressure, is a cardiovascular condition characterized by abnormally reduced blood pressure. Blood pressure is the force of blood pushing against the walls of the arteries as the heart pumps out blood and is ...
, together with weakness, dizziness, and periods of semi- or unconsciousness is also reported. If not treated carefully,
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
will spread, causing skin, subcutaneous tissue, and muscle to separate from healthy tissue and eventually slough with serous exudate. The slough may be superficial or deep, sometimes down to the bone.
Gangrene Gangrene is a type of tissue death caused by a lack of blood supply. Symptoms may include a change in skin color to red or black, numbness, swelling, pain, skin breakdown, and coolness. The feet and hands are most commonly affected. If the ga ...
and secondary infections commonly occur and can result in loss of digits and limbs. The fatality rate highly depends on the severity of the bites and some other factors. Deaths can be exceptional and probably occur in less than 15% of all untreated cases (usually in 2–4 days from complications following blood volume deficit and
disseminated intravascular coagulation Disseminated intravascular coagulation (DIC) is a condition in which blood clots form throughout the body, blocking Microvessel, small blood vessels. Symptoms may include chest pain, shortness of breath, leg pain, problems speaking, or problems ...
), although some reports show that severe envenomations have a 52% mortality rate. Most fatalities are associated with poor clinical management and neglect.


References


Further reading

*Access Professional Development. 2022. ''Puff Adder (Bitis arietans)''. nlineAvailable: https://accesspd.co.za/species/PuffAdder (Accessed: 02/02/2022) * Boulenger GA. 1896. ''Catalogue of the Snakes in the British Museum (Natural History). Volume III., Containing the...Viperidæ.'' London: Trustees of the British Museum (Natural History). (Taylor and Francis, printers.) xiv + 727 pp. + Plates I.- XXV. (''Bitis arietans'', pp. 493–495.) * Branch, Bill. 2004. ''Field Guide to Snakes and Other Reptiles of Southern Africa''. Third Revised edition, Second impression. Sanibel Island, Florida: Ralph Curtis Books. 399 pp. . (''Bitis arietans'', pp. 114–115 + Plates 3, 12.) * Broadley DG, Cock EV. 1975. ''Snakes of Rhodesia''. Zimbabwe: Longman Zimbabwe Ltd. 97 pp. *Broadley DG. 1990. ''FitzSimons' Snakes of Southern Africa''. Parklands (South Africa): J Ball & AD Donker Publishers. 387 pp. * Merrem B. 1820. ''Versuch eines Systems der Amphibien: Tentamen Systematis Amphibiorum.'' J.C. Krieger. Marburg. xv + 191 pp. + 1 plate.
(''"Vipera. Echidna. arietans"'', p. 152.) *Pienaar U de V. 1978. ''The reptile fauna of Kruger National Park''. National Parks Board of South Africa. 19 pp. *Sweeney RCH. 1961. ''Snakes of Nyasaland''. Zomba, Nyasaland: The Nyasaland Society and Nyasaland Government. 74 pp. *Turner RM. 1972. Snake bite treatment. Black Lechwe 10 (3): 24–33.


External links

* Accessed 9 December 2006. * . Accessed 1 March 2007.
Image of ''B. arietans'' bite that resulted in fasciotomy
a
South African Vaccine Producers
Accessed 26 July 2008.
Birds mob Puff Adder
– paper in ejournal Ornithological Observations {{Taxonbar, from=Q369667 Bitis Snakes of Africa Reptiles of North Africa Reptiles of Ethiopia Reptiles of Uganda Reptiles described in 1820 Taxa named by Blasius Merrem Articles containing video clips Snakes of Arabia