The black mamba (Dendroaspis polylepis) is a species of extremely venomous snake, a member of the family Elapidae native to parts of Sub-Saharan Africa. First formally described by Albert Günther in 1864, it is the second-longest venomous snake after the king cobra; mature specimens generally exceed 2 metres (6.6 feet) and commonly grow to 3 m (10 ft). Specimens of 4.3 to 4.5 m (14.1 to 14.8 ft) have been reported. Its skin colour varies from grey to dark brown. Juvenile black mambas tend to be paler than adults and darken with age.
The species is both terrestrial (ground-living) and arboreal (tree-living); it inhabits savannah, woodland, rocky slopes and in some regions, dense forest. It is diurnal and is known to prey on birds and small mammals. Over suitable surfaces, it can move at speeds up to 16 km/h (10 mph) for short distances. Adult black mambas have few natural predators.
In a threat display, the black mamba usually opens its inky-black mouth, spreads its narrow neck-flap and sometimes hisses. It is capable of striking at considerable range and may deliver a series of bites in rapid succession. Its venom is primarily composed of neurotoxins that often induce symptoms within ten minutes, and is often fatal unless antivenom is administered. Despite its reputation as a formidable and highly aggressive species, the black mamba only attacks humans if it is threatened or cornered. It is rated as least concern on the International Union for Conservation of Nature (IUCN)'s Red List of Threatened Species.
Although the black mamba had been known to missionaries and residents before 1860, the first formal description was made in 1864 by German-born British zoologist Albert Günther, who spelt the name Dendraspis polylepis. A single specimen was one of many species of snake collected by John Kirk, a naturalist who accompanied David Livingstone on the 1858–1864 Second Zambesi expedition. This specimen is the holotype and his housed in the Natural History Museum, London. The species' generic name is derived from the Ancient Greek words dendron (δένδρον), "tree", and aspis (ἀσπίς) "asp", and the specific epithet polylepis is derived from the Ancient Greek poly (πολύ) meaning "many" and lepis (λεπίς) meaning "scale". The term "mamba" is derived from the Zulu word "imamba". In Tanzania, a local Ngindo name is ndemalunyayo ("grass-cutter") because it supposedly clips grass.
In 1873, German naturalist Wilhelm Peters described Dendraspis Antinorii from a specimen in the museum of Genoa that had been collected by Italian explorer Orazio Antinori in what is now northern Eritrea. This was subsequently regarded as a subspecies and is no longer held to be distinct. In 1896, Belgian-British zoologist George Albert Boulenger combined the species Dendroaspis polylepis as a whole with the eastern green mamba (Dendroaspis angusticeps), a lumping diagnosis that remained in force until 1946 when South African herpetologist Vivian FitzSimons again split them into separate species. A 2016 genetic analysis showed the black and eastern green mambas are each others' closest relatives, and are more distantly related to Jameson's mamba (Dendroaspis jamesoni).
The black mamba is a long, slender, cylindrical snake with a "coffin-shaped" head with a somewhat pronounced brow ridge and a medium-sized eye. The adult snake's length typically ranges from 2 to 3 m (6 ft 7 in to 9 ft 10 in) but specimens have grown to lengths of 4.3 to 4.5 m (14.1 to 14.8 ft). It is the second-longest venomous snake species, exceeded in length only by the king cobra. The black mamba is a proteroglyphous (front-fanged) snake, with fangs up to 6.5 mm (0.26 in) in length, located at the front of the maxilla. The tail of the species is long and thin, the caudal vertebrae making up 17–25% of its body length. Black mambas weigh about 1.6 kg (3.5 lb) on average.
Specimens vary considerably in colour between olive, yellowish-brown, khaki and gunmetal but are rarely black. The scales of some individuals may have a purplish glow. Individuals occasionally display dark mottling towards the posterior, which may appear in the form of diagonal crossbands. Black mambas have greyish-white underbellies and the inside of the mouth is dark bluish-grey to nearly black. Mamba eyes range between greyish-brown to shades of black; the pupil is surrounded by a silvery-white or yellow colour. Juvenile snakes are lighter in colour than adults; these are typically grey or olive green and darken as they age.
Distribution and habitat
The black mamba inhabits a wide range within sub-Saharan Africa; it has been observed in north-east Democratic Republic of the Congo, south-western Sudan to Ethiopia, Eritrea, Somalia, Kenya, eastern Uganda, Tanzania, Burundi, Rwanda, and south to Mozambique, Swaziland, Malawi, Zambia, Zimbabwe and Botswana to KwaZulu-Natal in South Africa, and Namibia; then north-east across Angola to south-eastern Democratic Republic of the Congo.
The black mamba's distribution contains gaps within the Central African Republic, Chad, Nigeria and Mali. In 1954 the black mamba was recorded in the Dakar region of Senegal. This observation, and a subsequent observation that identified a second specimen in the region in 1956, has not been confirmed and thus the snake's distribution in this area is inconclusive.
The species prefers moderately dry environments such as light woodland and scrub, rocky outcrops and semi-arid savanna. It also inhabits moist savanna and lowland forests. It is not commonly found at altitudes above 1,000 m (3,300 ft), although its distribution does include locations at 1,800 m (5,900 ft) in Kenya and 1,650 m (5,410 ft) in Zambia. It is rated as a species of least concern on the International Union for Conservation of Nature (IUCN)'s Red List of Endangered species, based on its huge range across sub-Saharan Africa and no documented decline.
The black mamba is both terrestrial and arboreal. On the ground, it moves with its head and neck raised, and typically uses termite mounds, abandoned burrows, rock crevices and tree cracks as shelter. Black mambas are diurnal; in South Africa, they are recorded to bask between 7–10 am and again from 2–4 pm. They may return daily to the same basking site.
Skittish and often unpredictable, the black mamba is agile and can move quickly. When it perceives a threat, it retreats into brush or a hole. In the wild, black mambas seldom tolerate humans approaching more closely than about 40 metres (130 ft). When confronted it is likely to engage in a threat display, gaping to expose its black mouth and flicking its tongue. It also is likely to hiss and spread its neck-flap.
During the threat display, any sudden movement by the intruder may provoke the snake into performing a series of rapid strikes, leading to severe envenomation. The size of the black mamba and its ability to raise its head a large distance from the ground enable it to launch as much as 40% of its body length upwards, so mamba bites to humans may occur on the upper body. The black mamba's reputation for being ready to attack is exaggerated; it is usually provoked by perceived threats such as the blocking of its movements and ability to retreat. The species' reputed speed has also been exaggerated; it cannot move more quickly than 16 km/h (10 mph).
Reproduction and lifespan
The black mamba's breeding season takes place between September and February, following the drop in temperature which occurs from April to June. Rival males compete by wrestling, attempting to subdue each other by intertwining their bodies and wrestling with their necks. Some observers have mistaken this for courtship. During mating, the male will slither over the dorsal side of the female while flicking its tongue. The female will signal its readiness to mate by lifting its tail and staying still. The male will then coil itself around the posterior end of the female and align its tail ventrolaterally with the female's. Intermission may last longer than two hours and the pair remain motionless apart from occasional spasms from the male.
The black mamba is oviparous; the female lays a clutch of 6–17 eggs. The eggs are oval-shaped and elongated, measuring 60–80 mm (2.4–3.1 in) long and 30–36 mm (1.2–1.4 in) in diameter. When hatched, the young range from 40–60 cm (16–24 in) in length. They may grow quickly, reaching 2 m (6 ft 7 in) after their first year. Juvenile black mambas are very apprehensive and can be deadly like the adults. The black mamba is recorded to live up to 11 years and may live longer.
The black mamba usually hunts from a permanent lair, to which it will regularly return if there is no disturbance. It mostly preys on birds, particularly nestlings and fledglings, and small mammals like rodents, bats, hyraxes and bushbabies. They generally prefer warm-blooded prey but will also consume other snakes. The black mamba does not typically hold onto its prey after biting; rather it releases its quarry and waits for it to succumb to paralysis and death. The snake's potent digestive system has been recorded to fully digest prey in eight to ten hours.
Adult mambas have few natural predators aside from birds of prey. Young snakes have been recorded as prey of the Cape file snake. Mongooses, which have some resistance to mamba venom and are often quick enough to evade a bite, will sometimes take a black mamba for prey. The similarly predatory honey badger is also resistant. The mechanism in both mammals is thought to be that their muscular nicotinic acetylcholine receptors do not bind snake alpha-neurotoxins.
The black mamba is the most feared snake in Africa because of its size, aggression, toxicity and speed of onset of symptoms, and is classified as a snake of medical importance by the World Health Organization.[a] A survey of snakebites in South Africa from 1957 to 1963 recorded over 900 venomous snakebites, but only seven of these were confirmed black mamba bites; at that time no effective antivenom was widely available. Of the more than 900 bites, only 21 ended in fatalities, including all seven black mamba bites. The peak period for deaths is the species' breeding season from September to February, during which black mambas are most irritable. Bites are very rare outside Africa; snake handlers and enthusiasts are the usual victims.
Black mamba venom does not contain protease enzymes; its bites do not generally cause local swelling or necrosis, and a tingling sensation in the bitten area might be the only initial symptom. The snake tends to bite repeatedly and let go, so there might be multiple puncture wounds. Its bite can deliver about 100–120 mg of venom on average; the maximum recorded dose is 400 mg. The murine median lethal dose (LD50) when administered intravenously has been calculated at 0.32 and 0.33 mg/kg. Bites were very often fatal before antivenom was widely available.
The venom is predominantly neurotoxic; symptoms often become apparent within ten minutes. Early neurological symptoms that indicate a severe course include metallic taste, ptosis and gradual bulbar palsy. Other neurological symptoms include miosis, blurred or diminished vision, paresthesia, dysarthria, dysphagia, dyspnea, difficulty handling oral secretions, an absent gag reflex, fasciculations, ataxia, vertigo, drowsiness and loss of consciousness, and respiratory paralysis. Other more general symptoms include nausea and vomiting, abdominal pain, diarrhoea, sweating, salivation, goosebumps and red eyes. The bite of a black mamba can cause collapse in humans within 45 minutes or less. Without appropriate antivenom treatment, symptoms typically progress to respiratory failure, which leads to cardiovascular collapse and death. This typically occurs in 7 to 15 hours.
In 2015, the proteome (complete protein profile) of black mamba venom was assessed and published, revealing 41 distinct proteins and one nucleoside. The venom is composed of two main families of toxic agents, dendrotoxins (I and K) and (at a slightly lower proportion) three-finger toxins. Dendrotoxins are akin to kunitz-type protease inhibitors that interact with voltage-dependent potassium channels, stimulating acetylcholine and causing an excitatory effect, and are thought to cause symptoms such as sweating. Member of the three-finger family include alpha-neurotoxin, cardiotoxins, fasciculins and mambalgins. The most toxic components are the alpha-neurotoxins, which bind nicotinic acetylcholine receptors and hence block the action of acetylcholine at the postsynaptic membrane and cause neuromuscular blockade and hence paralysis. Fasciculins are anticholinesterase inhibitors that cause muscle fasciculation. The venom has little or no haemolytic, haemorrhagic or procoagulant activity. Mambalgins act as inhibitors for acid-sensing ion channels in the central and peripheral nervous system, causing a pain-inhibiting effect. There is research interest in their analgesic potential.
The composition of black mamba venom differs markedly from those of other mambas, all of which contain predominantly three-finger toxin agents. It is thought this may reflect the preferred prey items—small mammals for the mainly land-dwelling back mamba versus birds for the predominantly arboreal other mambas. Unlike many snake species, black mamba venom has little phospholipase A2 content.
Standard first aid treatment for any suspected bite from a venomous snake is the application of a pressure bandage to the bite site, minimisation of movement of the victim and conveyance to a hospital or clinic as quickly as possible. The lethal, neurotoxic nature of black mamba venom means an arterial tourniquet may be of benefit. Tetanus toxoid is given, though the main treatment is the administration of the appropriate antivenom. A polyvalent antivenom produced by the South African Institute for Medical Research is used to treat black mamba bites from many localities, and a new antivenom is being developed by the Universidad de Costa Rica's Instituto Clodomiro Picado.
Notable bite cases
Danie Pienaar, who was at various times from at least 2009 to 2017 head of South African National Parks Scientific Services and acting managing executive, survived the bite of a black mamba without antivenom in 1998. Despite the hospital physicians having declared it a "moderate" envenomation, Pienaar lapsed into a coma at one point and his prognosis was declared "poor". Upon arrival at hospital Pienaar was immediately intubated and placed on life support for three days. He was released from hospital on the fifth day. Remaining calm after being bitten increased his chances of survival, as did the application of a tourniquet.
In March 2008, 28-year-old British trainee safari guide Nathan Layton was bitten by a black mamba that had been found near his classroom at the Southern African Wildlife College in Hoedspruit. Layton was bitten by the snake on his index finger while it was being put into a jar, and first aid-trained staff who examined him determined he could carry on with lectures. He thought the snake had only brushed his hand. Layton complained of blurred vision either 20 minutes, 30 minutes, or an hour after being bitten, and collapsed shortly thereafter. He suffered cardiac arrest on the scene, and a post mortem identified the cause of death as snake bite.
American professional photographer Mark Laita was bitten on the leg by a black mamba during a photo-shoot of a black mamba at a facility in Central America. Bleeding profusely, he did not seek medical attention, and except for intense pain and local swelling overnight, he was not affected. This led him to believe that either the snake gave him a "dry bite" (a bite without injecting venom) or the heavy bleeding pushed the venom out. Some commenters on the story suggested that it was a venomoid snake (in which the venom glands are surgically removed), but Laita responded that it was not. Only later did Laita find that he had captured the snake biting his leg in a photograph.
In 2016, Kenyan woman Cheposait Adomo was attacked by three black mambas, one of which bit her repeatedly on the leg, in West Pokot County, Kenya. People coming to her aid drove off the other snakes, hacking two with a machete. After an attempt using traditional medicine, they placed her on a motorcycle and conveyed her 45 minutes to the nearest hospital, which had antivenom. She survived. Health services in Africa are trying to educate people about snakebite, as use of traditional medicine can delay vital treatment.
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