Black mamba

Dendroaspis polylepis also known as the black mamba, common black mamba, or the black-mouthed mamba is a species of large, highly venomous snake belonging to the Elapidae family and is native to Africa. The black mamba is the longest snake species in Africa, measuring between 2.5 and 3.2 m (8.2 and 10 ft) in length on average, and capable of growing to lengths of 4.4 m (14.6 ft). This species is named for the black colouration inside the mouth rather than the colour of its scales which varies from dull yellowish-green to a gun-metal grey. It is the fastest snake in the world, capable of moving at 4.32 to 5.4 metres per second (16–20 km/h, 10–12 mph). Black mambas are fast, nervous, lethally venomous, and when threatened, highly aggressive. They have been blamed for numerous human deaths, and African myths exaggerate their capabilities to legendary proportions. For these reasons, the black mamba is widely considered to be the world’s deadliest snake. Before the advent of black mamba antivenom, a bite from this species was 100% fatal, usually within about 20 minutes in very severe cases of envenomation.

Taxonomy and etymology
The black mamba is one of four species in the African snake genus Dendroaspis that are known as mambas. The species was first described in 1864 by Albert Günther, a German-born British zoologist, ichthyologist, and herpetologist. Soon after, a subspecies was identified, Dendroaspis polylepis antinorii (Peters, 1873), but this is no longer accepted as distinct. The genus and species name are derived from Ancient Greek words - Dendroaspis meaning "tree asp" (dendro is "tree", while aspis is "asp" which is understood to mean a "venomous snake") and polylepis, "many scaled", from poly "many" and lepis "scales". The name "black mamba" is given to the snake not because of its body colour but because of the ink-black colouration of the inside of its mouth, which it displays when threatened. In 1896, zoologist George Albert 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.

Physical Description
The adult black mamba's back skin color is olive, brownish, gray, or sometimes khaki in color. A young snake is lighter in colour, appearing gray or olive green, but not light enough to be confused with the different species of green mamba. Its underbody is cream-colored, sometimes blended with green or yellow. Dark spots or blotches may speckle the back half of the body, and some individuals have alternating dark and light scales near the posterior, giving the impression of lateral bars. The inside of the mouth is a dark blue to inky black in colour. The head is large but narrow and elongated, with a distinct shape of a coffin. It is a proteroglyphous snake, meaning it has immovable, fixed fangs at the front of the maxilla. The eyes are dark brown to black, with a silvery-white to yellow edge on the pupils. These snakes are strong but slender in body: adult specimens are 2.5 to 3 m (8.2 to 9.8 ft) in length on average, but specimens measuring 3.1 to 3.75 m (10 to 12.3 ft) are relatively common, and some specimens have reached lengths of 4.4 m (14.6 ft). The longest scientifically measured, wild-caught black mamba recorded was 4.48 metres (14.7 ft) long and weighed 11.7 kilograms (26 lb), found in Zimbabwe. Black mambas weigh about 5.6 kilograms (12 lb) on average, though they can weigh up to 12.5 kilograms (28 lb). There is no real sexual dimorphism, and both male and female snakes of this species have a similar appearance and tend to be similar in size. The species is the second-longest venomous snake in the world, exceeded in length only by the King cobra (Ophiophagus hannah). Information regarding the lifespan of snakes in the wild is sparse; the longest recorded lifespan of a captive black mamba is 14 years, but actual maximum lifespans could be much greater. As black mambas age, their colouration tends to get darker.

Scalation
There are 23-25 rows of smooth dorsal scales at midbody (rarely 21 rows), 248-281 ventrals and 109-132 paired subcaudals. The anal shield is divided. There are 7-10 upper labials, with the fourth (or third and fourth) entering the eye, and 11-13 lower labials (sometimes 10 or 14). There are three (sometimes four) preoculars and three or four (sometimes two or five) postoculars. Temporals are variable, usually 2+3.

Geographical Distribution
Although it is a large diurnal snake, the distribution of the black mamba is the subject of much confusion in research literature, indicating the poor status of African herpetological zoogeography. However, the distribution of the black mamba in eastern Africa and southern Africa is well documented. Pitman (1974) gives the following range for the species' total distribution in Africa: northeastern Democratic Republic of the Congo, southwestern Sudan, South Sudan to Ethiopia, Eritrea, Somalia, southern Kenya, eastern Uganda, throughout Tanzania, southwards to Mozambique, Swaziland, Malawi, Zambia, Zimbabwe and Botswana to KwaZulu-Natal in South Africa, and into Namibia; then northeasterly through Angola to the southeastern part of the Democratic Republic of Congo. According to CITES, the species is also found in Lesotho, Rwanda and Djibouti. The black mamba is not commonly found above altitudes of 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. West of Ethiopia, it has a curious distribution, with few records. There is a single record from the Central African Republic, two from Burkina Faso, and two unconfirmed sightings from Senegal, one from the Gambia, and a possible sighting in Cameroon. These sightings may indicate improper documentation, remaining populations from what was once a larger range, or new populations, indicating a growing range. The black mamba was 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, have not been confirmed and thus the species' distribution in West Africa is inconclusive. The black mamba's western 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.

Habitat
The black mamba has adapted to a wide variety of climates, ranging from savanna, woodlands, farmlands, rocky slopes, dense forests and humid swamps. The grassland and savanna woodland/shrubs that extend all the way from southern and eastern Africa to central and western Africa are the black mamba's typical habitat. The snake prefers more arid environments, such as semiarid, dry bush country, light woodland, and rocky outcrops. This species likes areas with numerous hills, as well as riverine forests. Black mambas often make use of abandoned termite mounds and hollow trees for shelter. The abandoned termite mounds are especially used when the snake is looking for somewhere to cool off, as the mounds are sort of a "natural air-conditioning" system. The structure of these mounds is very complex and elaborate. They have a network of holes, ducts, and chimneys that allow air to circulate freely, drawing heat away from the nest during the day - though without taking too much valuable moisture - while preventing the nest cooling too much at night. As a territorial species, though, the black mamba will always return to its territory or lair if left undisturbed.

Conservation status
This species is classified as Least Concern (LC) on the IUCN Red List of Threatened Species (v3.1, 2011). The conservation status of this species was last assessed in 2010 and it was classed as such due to its very large distribution throughout sub-Saharan Africa. Besides its very large geographical distribution, the species has no specific threats or predators that have been reported and this species is not undergoing significant population declines.

Behaviour
Although its scientific name seems to be indicative of tree climbing, the black mamba is mainly a terrestrial and diurnal snake. However, it is also arboreal, but much less so than the African green mambas. This species is an excellent climber which is equally at home on the ground and in trees. It is a very alert, high-strung and agile snake. It is known to be capable of reaching speeds of around 20 kilometers per hour (12 mph), travelling with up to a third of its body raised off the ground. Over long distances, the black mamba can travel 11 to 19 kilometers per hour (6.8 to 12 mph), but it can reach a speed of 16 to 20 kilometers per hour (9.9 to 12 mph) in short bursts, and it has been recorded at speeds of 23 kilometers per hour (14 mph), making it the fastest terrestrial snake species in the world. The black mamba is a territorial snake, having a favoured home usually in an abandoned termite mound, a hollow tree or log, or a rock crevice. It will actively defend its territory very aggressively. Although it is a shy and secretive snake in general, it always seeks to escape when a confrontation occurs, unless cornered, in which case the black mamba can put up a fearsome display of defense and aggression. When cornered, it mimics a cobra by spreading a neck-flap; exposing its black mouth, it lifts up to a third of its body up off the ground, and hisses. If the attempt to scare away the attacker fails, it will strike repeatedly. Many snake experts have cited the black mamba as the world's most aggressive snake, noting tendency to actively attack without provocation (B. Johnson, July 13, 2000; Hunter, 1998). According to Swaziland-born snake handler and snake expert Thea Litschka-Koen:


 * "black mambas will kill a dog or several dogs if threatened and it happens quite often. We also find dead cows and horses! We were called by the frantic family late one evening. When we arrived minutes later, two small dogs had already died and two more were showing severe symptoms of envenomation. Within 15 minutes we had found and bagged the snake. By this time the other two dogs were also dead. The snake must have been moving through the garden when it was attacked by the dogs. It would have struck out defensively, biting all the dogs that came within reach. The snake was bitten in several places on its body as well and died about a week later."

Communication and perception
Black mambas show little deviation from the common methods of communication and perception found in snakes. They use their eyesight mainly for detection of motion, and sudden movements will cause them to strike. The tongue is extended from the mouth to collect particles of air, which are then deposited in the vomeronasal organ on the roof of the mouth, which acts as a chemosensory organ. They have no external ears, but are quite adept at detecting vibrations from the ground. Like many snakes, when threatened, they will display aggression with a set of signals warning of the possibility of attack.

Hunting and prey
When hunting, the black mamba is often seen travelling with its head raised well above ground level, quickly moving forward in search of prey. Once prey is detected, the black mamba "freezes" before hurling itself forward and issuing several quick bites, swiftly killing its prey. If the prey attempts to escape, the black mamba will follow up its initial bite with a series of strikes. It will release larger prey after biting it, but smaller prey, such as birds or rats, are held until the prey's muscles stop moving. Black mambas feed on a variety of prey, especially mammals, including hyraxes, rats, mice, squirrels, bats, bushbabies and elephant shrews. They have also been known to prey on birds and small chickens, as well as other snakes. A large specimen has even been recorded eating a young blue duiker (Philantomba monticola). After ingestion, powerful acids digest the prey, sometimes within eight to 10 hours.

Reproduction
Black mambas breed only once a year. The breeding season begins in the spring, which occurs around the month of September in the African regions where these snakes occur, as much of sub-Saharan Africa is in the Southern Hemisphere. In this period, the males fight over females. Agonistic behaviour for black mambas involves wrestling matches in which opponents attempt to pin each other’s head repeatedly to the ground. Fights normally last a few minutes, but can extend to over an hour. The purpose of fighting is to secure mating rights to receptive females nearby during the breeding season. Beyond mating, males and females do not interact. Males locate a suitable female by following a scent trail. Upon finding his mate, he will thoroughly inspect her by flicking his forked tongue across her entire body. Males are equipped with two hemipenes. After a successful and prolonged copulation, the eggs develop in the female’s body for about 60 days. During this period, the female seeks a suitable place to lay the eggs. Females prefer using abandoned termite mounds as nests. Mature females lay between 15 and 25 eggs, which they hide very well and guard very aggressively. The eggs incubate for about 60 days before hatching. The hatchlings are about 50 centimetres (20 in) in length and are totally independent after leaving the eggs, hunting and fending for themselves from birth. Young hatchlings are as venomous as the adults, but do not deliver as much venom per bite as an adult snake would.

Venom
The venom of the black mamba is virulently toxic. Being a protein of low molecular weight, the venom of the black mamba is extremely rapid-acting. The venom consists mainly of highly potent neurotoxins, cardotoxins, fasciculins, and calciseptine. Subcutaneous LD50 values for this species' venom varies greatly. Ernst and Zug et al. 1996 gave it a value of 0.05 mg/kg, while Stephen Spawls and Bill Branch list it as 0.28 mg/kg and Engelmann & Obst (1981) list it as 0.32 mg/kg.