Dendroaspis Angusticeps

The eastern green mamba (''Dendroaspis angusticeps'') is a highly venomous snake species of the mamba genus ''Dendroaspis'' native to the coastal regions of southern East Africa. Described by Scottish surgeon and zoologist Andrew Smith in 1849, it has a slender build with a bright green back and green-yellow ventral scales. Adult females average around in length, and males are slightly smaller.

A shy and elusive species, the eastern green mamba is rarely seen. This elusiveness is usually attributed to its arboreal habitat and green colouration, which acts as camouflage in its natural environment. It has also been observed to use ambush predation, like many vipers, contrary to the active foraging style typical of other elapid snakes. It preys on birds, eggs, bats, and rodents such as mice, rats, and gerbils.

Its venom consists of both neurotoxins and cardiotoxins. Symptoms of envenomation include swelling of the injection site, dizziness, and nausea, accompanied by difficulty breathing and swallowing, irregular heartbeat, and convulsions progressing to respiratory paralysis. Bites that result in severe envenomation can quickly be fatal.

Taxonomy

The eastern green mamba was first described as ''Naja angusticeps'' by Andrew Smith, a Scottish surgeon and zoologist, in 1849, who reported it from Natal and east to Maputo Bay. The specific name ''angusticeps'' is derived from the Latin word ''angustus'', "narrow", and ''ceps'', an abbreviated form of ''caput'' ("head") when used in a compound word. The German-British zoologist Albert Günther described ''Dendroaspis intermedius'' from the Zambezi River in northern Mozambique in 1865. This was subsequently synonymised with ''D. angusticeps''.

In 1896, the Belgian-British zoologist George Albert Boulenger combined the species ''Dendroaspis angusticeps'' with the black mamba (''Dendroaspis polylepis''), a lumping diagnosis that remained in force until 1946, when the South African herpetologist Vivian FitzSimons published a paper after examining approximately 50 eastern green mamba and 85 black mamba specimens. He concluded that the differences in build, scalation, colouration and behaviour warranted splitting them into separate species. The British biologist Arthur Loveridge augmented FitzSimons' work with material from outside South Africa, noting some overlap in scalation but supporting the separation. A 2016 genetic analysis showed the eastern green and black mambas are each others' closest relatives, their common ancestor diverging from a lineage that gave rise to Jameson's mamba (''Dendroaspis jamesoni'') and the western green mamba (''Dendroaspis viridis''), as shown in the cladogram below.

As well as being called the eastern green mamba, the species is also frequently known as the common green mamba, East African green mamba, white-mouthed mamba, or simply the green mamba.

Description

The eastern green mamba is a large snake, with a slightly compressed and very slender body with a medium to long tapering tail. Adult males average around in total length, while females average . This species rarely exceeds lengths of . In general, the total length is 4–4.3 times the length of the tail. The adult eastern green mamba has bright green upperparts—occasionally with isolated yellow scales—and a pale yellow-green belly. Sometimes they are duller-coloured before moulting. Juveniles are blue-green, becoming bright green when they are around long. The coffin-shaped head is long and slender, with a prominent canthus which is slightly demarcated from the neck. When threatened or otherwise aroused, the eastern green mamba is capable of flattening its neck area into a slight hood. The medium-sized eyes have round pupils, the borders of which have a narrow golden or ochre edge; the irises are olive green, becoming bright green posteriorly. The inside of the mouth may be white or bluish-white.

Other green snakes are often called "green mambas" in Southern Africa, including green forms of the boomslang (''Dispholidus typus''), which can be distinguished by their larger eyes and shorter heads. They are also venomous. Green bushsnakes of the genus '' Philothamnus'' are also commonly confused with smaller Eastern green mambas.

Scalation

The number and pattern of scales on a snake's body are a key element of identification to species level. The eastern green mamba has between 17 and 21 rows of dorsal scales at midbody, 201 to 232 ventral scales, 99 to 126 divided subcaudal scales, and a divided anal scale. Its mouth is lined with 7–9 supralabial scales above, the fourth one located under the eye, and 9–11 sublabial scales below. Its eyes have three preocular and 3–5 postocular scales.

Distribution and habitat

The eastern green mamba is native to regions near the coastlines of Southern Africa and East Africa. Its range extends from Kenya south through Tanzania, Malawi, and eastern Zimbabwe; it can also be found in Zanzibar and northern Mozambique. An isolated and genetically distinct population is found in South Africa from the extreme northeastern part of Eastern Cape along the KwaZulu-Natal coastline and into southern Mozambique.

An elusive species, it is primarily arboreal (living in trees) and usually well camouflaged in foliage. Some herpetologists believe its habitat is limited to tropical rainforests in coastal lowlands. Other experts believe it can also be found in coastal bush, and dune and montane forest. Unlike its close relative the black mamba, the eastern green mamba is rarely found in open terrain and prefers relatively dense, well-shaded vegetation. As well as wild forest habitats, it is also commonly found in thickets and farm trees such as citrus, mango, coconut, and cashew. In coastal East Africa, it is known to enter houses and may even shelter in thatched-roof dwellings. Specimens have been found at elevations up to above sea level.

Conservation status

The species' conservation status has not been assessed by the International Union for Conservation of Nature (IUCN). It is fairly common throughout its range, and populations are believed to be stable. Large concentrations of two to three individuals per hectare have been documented in coastal Kenya and southern Tanzania, and in one instance a group of five were seen in a single tree. Although populations are stable overall, habitat destruction and deforestation may pose a threat. In South Africa, it is rated as "vulnerable" as its habitat had become highly fragmented by coastal housing development.

Behaviour and ecology

A diurnal species, the eastern green mamba is active by day and sleeps at night coiled up in foliage or rarely a hollow within the trunk or branch. An agile snake and an adept climber, it is not commonly found on the ground though may come down to bask in the morning sun (thermoregulation). A 27-day study of the movement patterns of two adults found their activity area to be relatively small, comparable to other predators who ambush prey rather than hunt (in contrast to most elapid species, including other mambas, who tend to hunt and forage). The study's preliminary evidence shed light on the species' feeding methods, suggesting it may be primarily an ambush predator due to its sit-and-wait behaviour. This evidence does not preclude active foraging, however; a specimen was also observed systematically hunting a sleeping bat.

There is no evidence that the eastern green mamba migrates. Thought to be relatively sedentary, it can remain in the same location for days, apparently moving most commonly to find food or mates. On average, individuals move only about per day. They generally avoid contact with people and other predators, attacking them only if they feel threatened.

Reproduction and lifespan

The eastern green mamba is solitary except during breeding season. Gravid females tend to be sedentary, but males actively seek and court prospective mates during the rainy season between April and June. Males have been seen engaging in agonistic behaviour; they may fight each other over mating opportunities, or possibly to establish a dominance hierarchy. Typically, one male initiates a fight by moving on top of the other's body and tongue-flicking, after which the two entwine and push, attempt to pin the other's head to the ground. Male combat can last for several hours, but is less aggressive than that commonly seen in the black mamba; combatants do not bite each other.

Males locate females by following a scent trail. The male courts the female by aligning his body along the female's while rapidly tongue-flicking. If the female is receptive to mating, she lifts her tail; cloacal juxtaposition follows. Courtship and mating take place in trees, after which the female can lay 4–17 eggs (10–15 on average), occurring in October and November. The eggs are small and elongated, usually 47–58 x 25–28 mm, and usually laid in leaf litter in hollow trees. The incubation period is around three months. When the young emerge, they are approximately in the wild, and in captivity. They usually reach adult colouration at . Hatchlings tend to grow to in their first year. As they age, their growth rates decrease but never completely stop.

The oldest recorded eastern green mamba was a captive specimen that lived 18.8 years. Another captive specimen lived 14 years.

Diet

The eastern green mamba preys primarily on birds and their eggs, and small mammals including bats. It is also believed to eat arboreal lizards. It uses a sit-and-wait strategy, though one specimen was recorded actively hunting sleeping bats. The species is also known to raid the nests of young birds. Sit-and-wait tactics may be successful with highly mobile prey, such as adult birds or rodents. Documented prey include the sombre greenbul, which occur in dense areas of natural and cultivated vegetation along Kenya's coastline. Ionides and Pitman (1965) reported a large bushveld gerbil in the stomach of a green mamba in Tanzania. Although the bushveld gerbil does not occur in Kenya, green mambas prey on the seven species of gerbil that inhabit parts of its range.

Predators

The eastern green mamba has few natural predators. Humans, mongooses, snake eagles and genets commonly prey on it, and hornbills and other snakes prey on juveniles.

Venom

The eastern green mamba has the least toxic venom of the three green mamba species, but it is still highly venomous. Although the most commonly encountered green mamba, it generally avoids people. The peak period for bites is the species' breeding season from September to February, during which they are most irritable. A survey in southern Africa from 1957–1979 recorded 2,553 venomous snakebites, 17 of which were confirmed as eastern green mambas. Of these 17, 10 had symptoms of systemic envenomation, though no victims died. The snake tends to bite repeatedly, and one bite can contain 60–95mg of venom by dry weight. The