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===Reproduction===
===Reproduction===
The eastern green mamba is [[Solitary animal|solitary]], except during breeding season, when they are most active. Although gravid females are relatively sedentary, males will actively search out and court females between April and June during the longer rainy season, and males have been documented to fight each other over potential mating opportunities, or possibly to establish a [[Dominance hierarchy|dominance hierarchy]]. One male initiates combat by moving on top of the other’s body and tongue-flicking, after which the two snakes “intertwine their necks and bodies, and push against each other” These combats may last for several hours, but combat between males of this species don't ever include biting and the nature of the combat is never as aggressive and/or vicious as commonly seen among the eastern green mamba's much larger cousin, the [[black mamba]]. Males find females by following a scent trail. The male initiates courtship by attempting to align its body along the female’s while rapidly tongue-flicking. If the female is receptive to [[mating]], she will lift her tail and cloacal juxtaposition will follow shortly. Courtship and mating take place in the trees, after which the female lays anywhere between 4-17 eggs (average of 10 to 15 eggs are laid), which occurs in the summer months of October and November.<ref name=H&M89/> The eggs are white and elongated, usually measuring 65x35 millimetres.<ref name="Broadley83"/> The eggs are usually laid in a hollow tree, among decaying vegetation, or leaf litter. The [[Egg incubation|incubation period]] is 10 to 12 weeks. When the young emerge out of the eggs, they are approximately {{convert|30|to|40|cm|in}}<ref name=S&B95/> or around an average of {{convert|44|cm|in}}<ref name=H&M89>{{cite journal|last=Haagner|first=GV|coauthors=Morgan, DR|title=The captive propagation of the Eastern green mamba Dendroaspis angusticeps|journal=International Zoo Yearbook|date=January, 1989|volume=28|issue=1|pages=195–199|doi=10.1111/j.1748-1090.1989.tb03280.x|accessdate=26 April 2014}}</ref> in length, and they're highly venomous right at birth. Individuals of this species usually reach adult coloration at a length of {{convert|60|to|75|cm|in}}<ref name="Broadley83"/><ref name=S&B95/> Hatchlings tend to grow {{convert|50|to|80|cm|in}} in length in the first year of life. As individuals age, their growth rates decrease but do not stop.<ref name=H&M89/>
The eastern green mamba is [[Solitary animal|solitary]], except during breeding season, when they are most active and males engage in combat and males and females mate. Gravid females tend to be sedentary, but males will actively search out and court females during the long rainy season, which is between the months of April and June. Males have been observed engaging in [[agnostic behaviour]] and may fight each other over potential mating opportunities, or possibly to establish a [[Dominance hierarchy|dominance hierarchy]]. Typically, a male initiates a fight by moving on top of the other’s body and tongue-flicking, after which the two snakes “intertwine their necks and bodies, and push against each other” in an attempt to pin each other's head repeatedly to the ground. Male-male combat can last for several hours, but combat between males of this species don't ever include biting and the nature of the combat is never as aggressive and/or vicious as commonly seen among the eastern green mamba's much larger cousin, the [[black mamba]]. 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. Depending on whether the female is receptive to [[mating]], she will lift her tail and cloacal juxtaposition will follow shortly. Courtship and mating take place in the trees, after which the female lays anywhere between 4-17 eggs (average of 10 to 15 eggs are laid), which occurs in the summer months of October and November.<ref name=H&M89/> The eggs are white and elongated, usually measuring 65x35 millimetres.<ref name="Broadley83"/> The eggs are usually laid in a hollow tree, among decaying vegetation, or leaf litter. The [[Egg incubation|incubation period]] is 10 to 12 weeks. When the young emerge out of the eggs, they are approximately {{convert|30|to|40|cm|in}}<ref name=S&B95/> or around an average of {{convert|44|cm|in}}<ref name=H&M89>{{cite journal|last=Haagner|first=GV|coauthors=Morgan, DR|title=The captive propagation of the Eastern green mamba Dendroaspis angusticeps|journal=International Zoo Yearbook|date=January, 1989|volume=28|issue=1|pages=195–199|doi=10.1111/j.1748-1090.1989.tb03280.x|accessdate=26 April 2014}}</ref> in length, and they're highly venomous right at birth. Individuals of this species usually reach adult coloration at a length of {{convert|60|to|75|cm|in}}<ref name="Broadley83"/><ref name=S&B95/> Hatchlings tend to grow {{convert|50|to|80|cm|in}} in length in the first year of life. As the hatchlings age, their growth rates decrease but they never stop completely growing.<ref name=H&M89/>


====Lifespan====
====Lifespan====
The longest living eastern green mamba was a captive specimen which lived for 18.8 years.<ref>{{cite web|title=AnAge entry for Dendroaspis angusticeps|url=http://genomics.senescence.info/species/entry.php?species=Dendroaspis_angusticeps|work=AnAge:The Animal Ageing and Longevity Database|publisher=Human Ageing Genomic Resources|accessdate=26 April 2014}}</ref> Another captive specimen lived for 14 years.<ref name="BB88"/> However, while it is feasible that wild individuals may live that long, they are thought to have shorter lifespans in general due to the threats of predation, habitat loss, and other environmental factors.
The longest living eastern green mamba was a captive specimen which lived for 18.8 years.<ref>{{cite web|title=AnAge entry for Dendroaspis angusticeps|url=http://genomics.senescence.info/species/entry.php?species=Dendroaspis_angusticeps|work=AnAge:The Animal Ageing and Longevity Database|publisher=Human Ageing Genomic Resources|accessdate=26 April 2014}}</ref> Another captive specimen lived for 14 years.<ref name="BB88"/> However, while it may be possible for wild specimens to live that long, they are thought to have shorter lifespans in general due to the threats of predation, habitat loss, disease, and other biological and environmental factors.


==Distribution and habitat==
==Distribution and habitat==

Revision as of 23:28, 28 April 2014

Eastern green mamba
Scientific classification Edit this classification
Missing taxonomy template (fix): Dendroaspis angusticeps
Binomial name
Dendroaspis angusticeps
(A. Smith, 1849)[1]
Eastern green mamba geographic range
Synonyms[2]
  • Naja angusticeps
    Smith, 1849
  • Naja angusticeps
    Duméril & Bibron, 1854
  • Dendraphis angusticeps
    Günther, 1858
  • Dendrospis [sic] angusticeps
    Boulenger, 1897
  • Dendroaspis sjöstedti
    Lönnberg, 1910
  • Dendraspis [sic] angusticeps
    Sternfeld, 1910
  • Dendroaspis angusticeps
    Razetti & Msuya, 2002

The eastern green mamba (Dendroaspis angusticeps), also known as the common mamba, East African green mamba, green mamba, or white-mouthed mamba, is a large arboreal and highly venomous snake species of the mamba genus Dendroaspis. This species of mamba was first described by a Scottish surgeon and zoologist in 1849. This snake mostly inhabits the coastal regions of southern East Africa. It is a relatively large species of venomous snake, with adult females averaging approximately 2.0 metres (6.6 ft) in total length, while males are slightly smaller on average. It is an arboreal species that preys on adult birds, eggs, bats, and terrestrial rodents such as mice, rats, and gerbils. Eastern green mambas are somewhat unique among mambas, and elapids in general. They are shy and elusive snakes which are rarely seen. The scarcity of observations is usually attributed to this species' cryptic colouration and arboreal lifestyle. However, low activity levels is associated with a sedentary foraging pattern might be an alternative explanation for its inconspicuousness. A 27 day observation of two adult eastern green mambas from the coastal forest of Gede/Jimba, Kenya conducted by Michael Angilletta showed that the eastern green mamba is not a typical active foraging elapid, but rather it is a "sit-and-wait" or ambush type predator (like most Viperidae).

Like other species of mamba, the eastern green mamba is a highly venomous species, capable of killing several humans with a single bite. It's venom is made up of pre-synaptic and post-synaptic neurotoxins (dendrotoxins), cardiotoxins, calcicludine, and fasciculins. The venom itself is of low molecular weight and is extremely rapid-acting, spreading through tissue very quickly and effectively. The dendrotoxins common to all species within the genus Dendroaspis is the most rapid-acting snake venom toxin known, so although this species is not aggressive and is not a major cause of snakebite incidents in Africa, the mortality rate associated with those bitten is rather high. A bite rapidly progressess to severe, life-threatening symptoms that are hallmarks of mamba bites. These symptoms include swelling of the bite site, dizziness, and nausea, accompanied by difficulty breathing and swallowing, irregular heartbeat, convulsions, and rapid progression to respiratory paralysis. Bites that produce severe envenomation can be rapidly fatal. Case reports of rapidly fatal outcomes, in as little as 30 minutes, have been recorded for this species.

Etymology

The eastern green mamba is classified under the genus Dendroaspis of the family Elapidae. Dendroaspis angusticeps was first described by a Scottish surgeon and zoologist, Dr. Andrew Smith in 1849.[3] The generic name, Dendroaspis, is derived from Ancient Greek words – Dendro, which means "tree",[4] and aspis (ασπίς) or "asp", which' is understood to mean "shield", but it also designate "cobra" or simply "snake". In old text, aspis or asp was used to refer to Naja haje (in reference with the hood, like a shield).[5] Thus, "Dendroaspis" literally means tree snake, which refers to the arboreal nature of most of the species within the genus. The specific name angusticeps is derived from the Latin word angustus, which means "narrow"[6] and -ceps is also Latin and is derived from the word "cephalicus" which means "head" or "of or relating to the head",[7] calling attention to the long narrow head of this species. In addition to being called the eastern green mamba, this species is also commonly known as the common green mamba, East African green mamba, white-mouthed mamba, or just simply the green mamba.[8]

Taxonomy

The eastern green mamba is classified under the genus Dendroaspis of the family Elapidae. The genus was first described by the German ornithologist and herpetologist Hermann Schlegel in 1848.[9] Slowinski et al (1997) pointed out that the relationships of the African genus Dendroaspis are problematical.[10] However, evidence suggests that Dendroaspis, Ophiophagus, Bungarus, and Hemibungarus form a solid non-coral snake Afro-Asiatic clade.[11]

Biology

Physical description

An eastern green mamba

The eastern green mamba is a fairly large, with a slightly compressed, and very slender bodied snake with a medium to moderately long tapering tail. Adult males average around 1.8 metres (5.9 ft) in total length, while females average 2.0 metres (6.6 ft) in total length. This species rarely exceeds lengths of 2.5 metres (8.2 ft). In general, the total length is 4-4.3 times the length of the tail.[12][13] The head is narrow, elongate, and coffin-shaped with a distinct canthus and slightly distinct from the neck. The neck may be flattened when snake is aroused, but there is no real hood. The eyes are medium in size with round pupils. This species has enlarged front fangs located at anterior end of maxillary bone (proteroglyphous) at the very front of the maxilla, which can rotate at its articulation with the prefrontal bone. However, there are no other solid teeth on the maxillary bone. At the front of the lower jaw is a pair of long, recurved, fang-like solid teeth, followed behind by a distinct interspace and numerous small teeth. Their eyes are medium in size with round pupils.[8][14]

Scalation

Dorsal scales are oblique, smooth and narrow. Coloration in this species is bright green dorsally and yellow-green ventrally, with a few bright yellow scales scattered on the flanks in some specimens. Juveniles are blue-green, and develop the brighter green adult coloration anteriorly to posteriorly in successive sheddings of the skin. Most individuals over 60 centimetres (24 in) in total length have the full adult coloration, but even some adults may return to a darker bluish green just before shedding. The border of the pupil may have a narrow bright ochre to golden yellow edge, and the posterior border of the iris may become bright green. The inside of the mouth may be white or bluish white. The males of this species usually have fewer ventral scales than the females.[8][12][13]

The head, body, and tail scalation of the eastern green mamba:[15]

Reproduction

The eastern green mamba is solitary, except during breeding season, when they are most active and males engage in combat and males and females mate. Gravid females tend to be sedentary, but males will actively search out and court females during the long rainy season, which is between the months of April and June. Males have been observed engaging in agnostic behaviour and may fight each other over potential mating opportunities, or possibly to establish a dominance hierarchy. Typically, a male initiates a fight by moving on top of the other’s body and tongue-flicking, after which the two snakes “intertwine their necks and bodies, and push against each other” in an attempt to pin each other's head repeatedly to the ground. Male-male combat can last for several hours, but combat between males of this species don't ever include biting and the nature of the combat is never as aggressive and/or vicious as commonly seen among the eastern green mamba's much larger cousin, the black mamba. 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. Depending on whether the female is receptive to mating, she will lift her tail and cloacal juxtaposition will follow shortly. Courtship and mating take place in the trees, after which the female lays anywhere between 4-17 eggs (average of 10 to 15 eggs are laid), which occurs in the summer months of October and November.[16] The eggs are white and elongated, usually measuring 65x35 millimetres.[12] The eggs are usually laid in a hollow tree, among decaying vegetation, or leaf litter. The incubation period is 10 to 12 weeks. When the young emerge out of the eggs, they are approximately 30 to 40 centimetres (12 to 16 in)[13] or around an average of 44 centimetres (17 in)[16] in length, and they're highly venomous right at birth. Individuals of this species usually reach adult coloration at a length of 60 to 75 centimetres (24 to 30 in)[12][13] Hatchlings tend to grow 50 to 80 centimetres (20 to 31 in) in length in the first year of life. As the hatchlings age, their growth rates decrease but they never stop completely growing.[16]

Lifespan

The longest living eastern green mamba was a captive specimen which lived for 18.8 years.[17] Another captive specimen lived for 14 years.[18] However, while it may be possible for wild specimens to live that long, they are thought to have shorter lifespans in general due to the threats of predation, habitat loss, disease, and other biological and environmental factors.

Distribution and habitat

Geographic range

This species is indigenous to southern East Africa, mostly along the coastline. In particular, the eastern green mamba's range extends from Kenya south through Tanzania, Mozambique, Malawi, eastern Zimbabwe, into South Africa as far as southern Natal and northern Pondoland.[8][12]

Habitat

The eastern green mambas is primarily arboreal, and while it does occasionally venture to the ground, it is most commonly found camouflaged in bushes and trees. There are reports that this species is limited to tropical rainforests in coastal lowlands,[16] however, according to other experts, this species can also be found in coastal bush, and dune and montane forest.[18] Unlike its close relative the black mamba (D. polylepis), this species is rarely found in open habitats and seems to prefer relatively dense, well-shaded vegetation. In addition to wild forest habitats, this species has also been found in farm trees such as citrus, mango, coconut, and cashew. In coastal east Africa they are known to enter houses and may even shelter in thatched roof dwellings. Specimens of this species have been been found at elevations up to 1,500 metres (4,900 ft) above sea level.[13]

Conservation status

As of June 26, 2011, the conservation status of Dendroaspis angusticeps has not been assessed by the IUCN (IUCN 2011). However, this species is fairly common within its range, and populations are thought to be robust. 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 D. angusticeps was observed in a single tree. Although populations of this species are thought to remain healthy, habitat destruction and deforestation may threaten their further stability.[13]

Behaviour and ecology

Behavior

A green mamba at a German Serpentarium

The eastern green mamba is a diurnal, arboreal, and secretive species of snake, and it tends to spend most of its time above the ground in relatively dense brush, where it is well camouflaged.[13] This species is not commonly found on land unless motivated by thirst, prey, or the need to bask in the sun (thermoregulation).[19] It is an alert, nervous, excellent climber and extremely agile snake. It sleeps at night in a tree coiled up in leafy clumps rather than seeking a tree hollow (although sometimes found in them).[8] In a study of the movement patterns of two adult specimens of this species over a 27-day period, the researcher found that their activity range areas to be very low, comparable to other predators who ambush prey rather than actively hunt them. This is in contrast to most elapid species, including other mambas, who tend to actively hunt or forage for prey. The study's preliminary evidence sheds some light on this species' method of hunting prey and suggests that it may be an ambush predator due to the sit-and-wait behavior displayed. However, this evidence does not preclude active foraging by this species. A specimen systematically hunting a sleeping bat was observed by William York.[20] There is no evidence that the eastern green mamba migrates; in fact, this species is thought to be relatively sedentary. It can remain in the same location for days at a time, apparently moving most commonly to find food or mates. On average, individuals of this species move only about 5.4 metres (18 ft) per day.[16][20] Unlike its much larger cousin the Black mamba, this mamba is more shy and not as aggressive or fearsome. It will avoid confrontation with humans or any other potential predators when possible, and will rather rely on its camouflage, or flee, than alert a potential threat of its presence. They are fast snakes, capable of moving 7 mph. They don't always strike, but under continuous harassment and provocation and especially if cornered, they may suddenly strike repeatedly in quick succession, often leading to severe envenomation.[19]

Diet

This mamba preys primarily on adult birds, eggs and rodents. This species has also been documented to prey on bats. It is also believed that this species eats arboreal lizards as well, but this has not been documented.[12][13][18] The preliminary evidence suggests that this species displays a sit-and-wait strategy of foraging. However, this evidence does not preclude active foraging by this species. One witness observed a specimen systematically hunt sleeping bats (J. Ashe, pers. comm. 1991).[20] They have also been known to raid the nests of young birds.[21] Sit-and-wait tactics may be successful with highly mobile prey, such as adult birds or rodents. Documented prey include the Sombre Greenbul, which inhabits dense portions of natural and cultivated vegetation on the coast of Kenya. Ionides and Pitman (1965) reported a large Bushveld gerbil in the stomach of a green mamba in Tanzania. While this species does not extend into Kenya, it is assumed that green mambas will prey on any of the seven species of gerbil that inhabit various portions of its range.[20]

Predators

The eastern green mamba has a few natural predators. Humans, mongooses, snake eagles, and genets commonly prey on this species of mamba. Hornbills and other snakes tend to prey on juvenile green mambas.[20]

Venom

The eastern green mamba has a very rapid-acting venom

The eastern green mamba is an especially venomous snake. The venom consists of both pre-synaptic and postsynaptic neurotoxins (dendrotoxins), cardiotoxins, calcicludine, and fasciculins. The average venom yield per bite is 80 mg according to Engelmann and Obst (1981),[22] while Minton (1974) gives it a range of 60–95 mg (dry weight).[8] The subcutaneous LD50 is 1.3 mg/kg.[13] The LD50 in mice through the IV route is 0.45 mg/kg.[23] Like all other mamba species, the toxicity of individual specimens within the same species can vary greatly based on several factors including geographical region, age, seasonal variation, diet, and so on. Local swelling is variable and sometimes absent after mamba bites. However, patients bitten by the eastern green mamba develop swelling of the entire bitten limb and also show mild haemostatic disturbances (Warrell DA; MacKay et al. 1966). The rare cases of local tissue damage usually resulted from bites on the fingers or the use of a tight tourniquet.[24] This species has caused bites to humans and many of the bites attributed to this species have often resulted in fatalities. The mortality rate of untreated bites is unknown but is thought to be quite high.[8] Symptoms of envenomation by this species include swelling of the bite site, dizziness, and nausea, accompanied by difficulty breathing and swallowing, irregular heartbeat, convulsions, rapid progression to respiratory paralysis. Bites that produce severe envenomation can be rapidly fatal. Case reports of rapidly fatal outcomes, in as little as 30 minutes, have been recorded for this species.[13]

References

  1. ^ "Dendroaspis angusticeps". Integrated Taxonomic Information System. Retrieved 30 October 2012.
  2. ^ "Dendroaspis angusticeps (SMITH, 1849)". The Reptile Database. Retrieved 17 February 2012.
  3. ^ Smith, Andrew (1848). Illustrations of the zoology of South Africa, Reptilia. London: Smith, Elder and Co.
  4. ^ "dendro-". Collins English Dictionary - Complete & Unabridged 10th Edition. HarperCollins Publishers. Retrieved 4 March 2014.
  5. ^ "aspis, asp". Dictionary.com Unabridged. Random House. Retrieved 4 March 2014.
  6. ^ "Definition of "angustus"". Numen - The Latin Lexicon. http://latinlexicon.org/index.php. Retrieved 26 April 2014. {{cite web}}: External link in |publisher= (help)
  7. ^ "Definition of "-ceps"". Numen - The Latin Lexicon. http://latinlexicon.org/index.php. Retrieved 26 April 2014. {{cite web}}: External link in |publisher= (help)
  8. ^ a b c d e f g "Dendroaspis viridis". Clinical Toxinology Resource. University of Adelaide. Retrieved 26 April 2014. Cite error: The named reference "WCH" was defined multiple times with different content (see the help page).
  9. ^ "Dendroaspis". Integrated Taxonomic Information System. Retrieved 9 December 2013.
  10. ^ Slowinski, JB; Knight, A; Rooney, AP (December 1997). "Inferring species trees from gene trees: A phylogenetic analysis of the Elapidae (Serpentes) based on the amino acid sequences of venom proteins". Molecular Phylogenetics and Evolution. 8 (3): 349–62. doi:10.1006/mpev.1997.0434. PMID 9417893.
  11. ^ Castoe, TA. "Higher-level phylogeny of Asian and American coralsnakes, their placement within the Elapidae (Squamata), and the systematic affinities of the enigmatic Asian coralsnake Hemibungarus calligaster (Wiegmann, 1834)" (PDF). Zoological Journal of Linnean Society. 151: 809–831. Retrieved 26 April 2014. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  12. ^ a b c d e f Broadley, D. (1983). "9". In Fitzsimmons, VFM (ed.). Fitzsimmons' Snakes of Southern Africa (Reprint, revised ed.). Johannesburg, South Africa: Delta Books, LTD. ISBN 978-0-908387-04-5.
  13. ^ a b c d e f g h i j Spawls, S., Branch, B. (1995). The Dangerous Snakes of Africa. Blandford. pp. 49–51. ISBN 978-0-88359-029-4.{{cite book}}: CS1 maint: multiple names: authors list (link)
  14. ^ Zug, George R. (1996). Snakes in Question: The Smithsonian Answer Book. Washington D.C., USA: Smithsonian Institution Scholarly Press. ISBN 1-56098-648-4.
  15. ^ Marais, J (2004). A Complete Guide to Snakes of Southern Africa. Cape Town: Random House Struik Publishers. pp. 98–99. ISBN 978-1-8-6872-932-6.
  16. ^ a b c d e Haagner, GV (January, 1989). "The captive propagation of the Eastern green mamba Dendroaspis angusticeps". International Zoo Yearbook. 28 (1): 195–199. doi:10.1111/j.1748-1090.1989.tb03280.x. {{cite journal}}: |access-date= requires |url= (help); Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  17. ^ "AnAge entry for Dendroaspis angusticeps". AnAge:The Animal Ageing and Longevity Database. Human Ageing Genomic Resources. Retrieved 26 April 2014.
  18. ^ a b c Branch, B. (1988). Branch's Field Guide Snakes Reptiles Southern Africa. Curtis Publishing, Ralph. ISBN 978-0-88359-023-2.
  19. ^ a b O'Shea, Mark (12 September 2005). Venomous Snakes of the World. New Jersey, USA: Princeton University Press. ISBN 0-691-12436-1.
  20. ^ a b c d e Angilletta, MJ. (1994). "Sedentary behaviors by Green Mambas Dendroaspis angusticeps" (PDF). Herpetological Natural History. 2 (2): 105–111. Retrieved 26 April 2014.
  21. ^ Phelps, T. (1989). Poisonous Snakes. London: Blandford. p. 272. ISBN 978-0-71-372114-0.
  22. ^ Engelmann, Wolf-Eberhard (1981). Snakes: Biology, Behavior, and Relationship to Man. Leipzig; English version NY, USA: Leipzig Publishing; English version published by Exeter Books (1982). p. 51. ISBN 0-89673-110-3.
  23. ^ Department of the Navy Bureau of Medicine and Surgery (2013). Venomous Snakes of the World: A Manual for Use by U.S. Amphibious Forces. Skyhorse Publishing. p. 93. ISBN 978-1-62087-623-7.
  24. ^ "WHO - Guidelines for the Prevention and Clinical Management of Snakebite in Africa". WHO Regional Office for Africa. World Health Organization. Retrieved 21 December 2011.
  • Smith, A. 1849. Illustrations of the zoology of South Africa. Reptilia. Smith, Elder & Co. London.

External links

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