In red, range of D. polylepis
The black mamba (Dendroaspis polylepis) is a venomous snake endemic to sub-Saharan Africa. It takes its common name not from the colour of its scales, but from the interior of its mouth, which is inky-black. It is the longest species of venomous snake on the African continent, with a length typically ranging from 2 meters (6.6 ft) to 3 meters (9.8 ft) and up to 4.3 to 4.5 meters (14.1 to 14.8 ft). It is among the fastest-moving snakes in the world, capable of moving at 11 km/h (6.8 mph) over short distances.
Black mambas breed annually and mating occurs in the early spring. Females lay eggs which gestate over 80 to 90 days before hatching. Juvenile black mambas are lighter in colour than adults and darken with age. Although mambas are typically tree-dwelling snakes, the black mamba is only occasionally arboreal, preferring to build lairs in terrestrial habitats. The black mamba is found across a range of terrain from savannah, woodlands, rocky slopes and dense forests. It is diurnal and chiefly an ambush predator, known to prey on hyrax, bushbabies and other small mammals. Adult black mambas have few predators in the wild.
The venom of the black mamba is highly toxic, potentially causing collapse in humans within 45 minutes or less from a single bite. Without effective antivenom therapy, death typically occurs in 7–15 hours. The venom is chiefly composed of neurotoxins, specifically dendrotoxin. The black mamba is capable of striking at considerable range and occasionally may deliver a series of bites in rapid succession. Despite its reputation for being highly aggressive, like most snakes, it usually attempts to flee from humans unless threatened or cornered.
The black mamba is classified under the genus Dendroaspis of the family Elapidae and the species D. polylepis. The black mamba was first described in 1864 by Albert Günther, a German-born British zoologist, ichthyologist, and herpetologist. In 1873, Wilhelm Peters described two subspecies: the nominotypical D.polylepis polylepis and also D.polylepis antinorii. However, these are no longer held to be distinct. In 1896, Boulenger combined the species (Dendroaspis polylepis) as a whole with the eastern green mamba (Dendroaspis angusticeps) and they were considered a single species from 1896 until 1946, when FitzSimons split them into separate species again.
The generic name, Dendroaspis, is derived from Ancient Greek Dendro, which means "tree", and aspis or "asp", which is understood to mean "shield", but also denotes "cobra" or simply "snake". In ancient texts, aspis or asp was often used to refer to the Egyptian cobra (Naja haje), in reference to its shield-like hood. Thus, "Dendroaspis" literally means tree snake, which refers to the arboreal nature of most of the species within the genus. The specific name polylepis is derived from the Ancient Greek poly meaning "many" or "multiple" and lepis meaning "scale" giving the literal meaning "many scaled". Most likely in reference to the size and higher scale count of this species compared to other species in the genus.
Contrary to its common name, the black mamba is not actually black. It takes its name from the interior of the snake's mouth, which is inky-black in colour. It is a large, round-bodied, slender snake, with a progressively tapering tail, and is of markedly stockier build than its close relatives Dendroaspis angusticeps and Dendroaspis viridis. The head is often said to be "coffin-shaped" with a somewhat pronounced brow ridge and a medium-sized eye. The black mamba's skin colouration can vary between olive-brown to grey hues, or sometimes khaki and some individuals may display dark mottling towards the posterior that may form oblique bars. The underbody is often pale yellow or cream coloured and the eyes are dark brown to black with a silver or pale yellow corona surrounding the pupil. Juvenile snakes are lighter in colour than adults, typically grey or olive green in appearance and darken with age. It is a proteroglyphous snake, with fangs up to 6.5 millimeters (0.26 in) in length located at the front of the maxilla. The adult snake's length ranges from 2 meters (6.6 ft) to 3 meters (9.8 ft) routinely but, according to some sources, specimens have reached lengths of 4.3 to 4.5 meters (14.1 to 14.8 ft). Black mambas weigh about 1.6 kilograms (3.5 lb) on average. A specimen of 1.41 meters (4.6 ft) was found to have weighed 651.7 g (1.437 lb). The species is the second longest venomous snake in the world, exceeded in length only by the king cobra of India and Southeast Asia.
The head, body and tail scalation of the black mamba:
Distribution and habitat
The black mamba occurs across a wide and occasionally fragmented range of sub-Saharan Africa. Specifically, the black mamba's range has been observed as: north east Democratic Republic of the Congo, south western Sudan to Ethiopia, Eritrea, Somalia, Kenya, eastern Uganda, Tanzania, Burundi, Rwanda, southwards to Mozambique, Swaziland, Malawi, Zambia, Zimbabwe and Botswana to KwaZulu-Natal in South Africa, and Namibia; then north easterly through 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. These gaps may lead physicians to misidentify the black mamba and administer an ineffective antivenom.
The black mamba was also recorded in 1954 in West Africa, in the Dakar region of Senegal. However, 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 black mamba is not commonly found at altitudes above 1,000 metres (3,300 ft), although the distribution of black mamba does reach 1,800 metres (5,900 ft) in Kenya and 1,650 metres (5,410 ft) in Zambia.
The black mamba is primarily terrestrial in nature, although it has been observed to be occasionally arboreal; typically inhabiting neglected areas of scrub, termite mounds, abandoned burrows and rock crevices. It has adapted to a variety of terrain ranging from savannah and woodlands to rocky slopes and dense forests. The black mamba prefers more arid environments such as light woodland and scrub, rocky outcrops, and semi-arid dry savannah.
Behaviour and ecology
The black mamba is a graceful but skittish and often unpredictable snake, capable of moving quickly and with great agility. It is shy and secretive by nature and, like most snakes, will try to avoid danger where it can. In the wild, a black mamba will seldom allow a close approach (within 40 meters). When confronted it can display great truculence and will gape, in mimicry of a cobra, by spreading its neck-flap, exposing its black mouth and flicking its tongue. Sometimes this behaviour may be accompanied by audible hissing. Any sudden movement at this stage may provoke the mamba into a series of rapid strikes leading to severe envenomation. Also, due to its size, the black mamba is able to raise its head well off the ground and in striking may be able to throw as much as 40% of its body upwards. This allows for considerable striking range, including humans at chest height. The black mamba's reputed readiness to attack is often much exaggerated and is typically the result of an interference in its movements; whether intentional or otherwise.
The black mamba is the fastest moving snake found in Africa, and one of the fastest moving snakes in the world—perhaps the fastest. There have been many exaggerated stories concerning the black mamba's speed of movement across the ground, and the elongated slender body can create the impression that it is moving faster than it really is. These stories include the myth that it can outrun a galloping horse or a running human. On 23 April 1906, on the Serengeti Plains, an intentionally provoked and angry black mamba was recorded at a speed of 11 km/h (6.8 mph), over a distance of 43 m (141 ft). A black mamba would almost certainly not be able to exceed 16 km/h (9.9 mph) and it can only maintain such relatively high speeds for short distances.
The black mamba is diurnal and chiefly an ambush predator. Hunting is usually conducted from a permanent lair, which it will regularly return to providing it is not disturbed. When hunting, it has been known to raise a large portion of its body off the ground. The black mamba does not typically hold onto prey after biting, instead releasing its quarry and waiting for it to succumb to paralysis and die. If prey attempts to escape or defend itself, the black mamba will often follow up its initial bite with a rapid series of strikes to incapacitate and quickly kill its prey. The snake has been known to prey on hyrax and rock hyrax, bushbabies, and bats. The black mamba has a potent digestive system and has been observed to fully digest prey within eight to ten hours.
Not many predators challenge an adult black mamba although it does face a few threats such as birds of prey, particularly snake eagles. Although all species of snake eagle commonly prey on snakes, there are two species in particular that do so with high frequency, including preying on black mambas. These are the black-chested snake eagle (Circaetus pectoralis) and the brown snake eagle (Circaetus cinereus). The Cape file snake (Mehelya capensis), which is apparently immune to all African snake venoms and preys on other snakes including venomous ones, is a common predator of black mambas (up to a size it can swallow). Mongooses which are also partially immune to venom, and are often quick enough to evade a bite, will sometimes tackle a black mamba for prey. Humans do not usually consume black mambas, but they often kill them out of fear.
Black mambas breed annually and mating occurs in the early spring, with male mambas locating a female by following her scent trail. After finding a potential mate the male will inspect the female by flicking his tongue over her entire body. Males possess hemipenes. Like most snakes female mambas are both oviparous and iteroparous. Egg-laying typically occurs during the middle of summer and egg clutches range from 6 to 17 eggs, with gestation occurring over approximately 80 to 90 days. During the mating season rival males may engage in combat that involves twisting their bodies around each other and raising their heads high off the ground, in a bid to subdue their opponent. This may sometimes be mistaken for mating.
Black mambas are solitary in nature and do not interact beyond mating and male-to-male combat. Following incubation, juvenile mambas break though the egg shell with an egg tooth and are born with fully developed venom glands. Thus, they are capable of inflicting a potentially lethal bite minutes after birth. The egg yolk is absorbed into the juvenile snake's body as a source of nourishment that sustains the young snake following birth.
There is little information available concerning the lifespan of wild black mambas, but the longest surviving captive example had a recorded lifespan of 11 years. It is possible that wild snakes may live significantly longer than this.
The black mamba's venom is composed of neurotoxins (dendrotoxin) and cardiotoxins as well as other toxins such as fasciculins. In an experiment, the most abundant toxin found in black mamba venom was observed to be able to kill a mouse in as little as 4.5 minutes. Based on the murine median lethal dose (LD50) values, the black mamba's toxicity from all published sources is as follows:
- (SC) subcutaneous (most applicable to real bites): 0.32 mg/kg, 0.28 mg/kg.
- (IV) intravenous: 0.25 mg/kg, 0.011 mg/kg.
- (IP) intraperitoneal: 0.30 mg/kg (average), 0.941 mg/kg. 0.05 mg/kg (the last quote doesn't make it clear if is either intravenous or intraperitoneal).
Its bites can deliver about 100–120 mg of venom on average and the maximum dose recorded is 400 mg. It is reported that before antivenom was widely available, the mortality rate from a bite was nearly 100%. The bite of a black mamba can potentially cause collapse in humans within 45 minutes, or less. Without effective antivenom therapy, death typically occurs in 7–15 hours. Presently, there is a polyvalent antivenom produced by the South African Institute for Medical Research to treat black mamba bites from many localities.
A bite from a black mamba causes initial neurological and neuromuscular symptoms which may commonly include headache and a metallic taste in the mouth, which may be accompanied by a triad of paresthesias, profuse perspiration and salivation. Other symptoms may include ptosis and gradual bulbar palsy. Localised pain or numbness around the bite site is common but not typically severe; therefore, application of a tourniquet proximal to the bite site is feasible and may assist in slowing the onset of prominent neurotoxicity. Without appropriate treatment, symptoms typically progress to more severe reactions such as tachydysrhythmias and neurogenic shock, leading to death by asphyxiation, cardiovascular collapse, or respiratory failure.
Attacks on humans
The black mamba is popularly regarded as the most dangerous and feared snake in Africa; to South African locals the black mamba's bite is known as the "kiss of death". However, attacks on humans by black mambas are rare, as they usually try to avoid confrontation and their occurrence in highly populated areas is not very common compared with some other species. Additionally, the ocellated carpet viper is responsible for more human fatalities due to snakebite than all other African species combined. 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 a time when effective antivenom was not widely available. Out of more than 900 bites, only 21 ended in fatalities, including all seven black mamba bites.
Reported bite cases
In 1998, Danie Pienaar, now head of South African National Parks Scientific Services, survived the bite of a black mamba without antivenom. Although no antivenom was administered, Pienaar was in serious condition, despite the fact the hospital physicians declared it a "moderate" black mamba envenomation. At one point, Pienaar lapsed into a coma and his prognosis was declared "poor". Upon arrival at hospital Pienaar was immediately intubated, given supportive drug therapy, put on mechanical ventilation and was placed on life support for three days, until the toxins were flushed out of his system. He was released from hospital on the fifth day. Pienaar believes he survived for a number of reasons. In an article in Kruger Park Times he said: "Firstly, it was not my time to go." the article also stated "The fact that he stayed calmed and moved slowly definitely helped. The tourniquet was also essential."
In another case, 28-year-old British student Nathan Layton was bitten by a black mamba and died of a heart attack in less than an hour in March 2008. The black mamba had been found near a classroom at the Southern African Wildlife College in Hoedspruit, where Layton was training to be a safari guide. Layton was bitten by the snake on his index finger while it was being put into a jar, but he didn't realize he'd been bitten. He thought the snake had only brushed his hand. Approximately 30 minutes after being bitten Layton complained of blurred vision. He collapsed and died of a heart attack, nearly an hour after being bitten. Attempts to revive him failed and he was pronounced dead at the scene.
In 2013, in a rare case of survival without treatment, 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. The bite ruptured an artery in his calf, and he was gushing blood profusely. Laita did not go to the doctor or the hospital, and except for the swollen fang marks giving him intense pain during the night, he was not affected and was fine physically. This led him to believe that the snake either gave him a "dry bite" (meaning without injecting venom) or that the heavy bleeding pushed the venom out. Some commenters to the story suggested that it was a venomoid snake (in which the venom glands are surgically removed). Laita responded that it was not the case. Only later, Laita found that he had captured the snake biting his leg in a photograph.
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