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DOI10.47051/BJCI8462 Published April 20, 2020. Open access.

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Amazonian Bushmaster (Lachesis muta)

Reptiles of Ecuador | Serpentes | Viperidae | Lachesis muta

English common names: Amazonian Bushmaster, South American Bushmaster.

Spanish common names: Verrugosa amazónica, verrugosa del Oriente, motolo, yamunga (Ecuador); cascabel muda, cuaima, mapaná rayo, pudridora verrugosa, verrugoso (Colombia); shushupe, matabuey, cuanira, kempirona, macapé (Perú); cuaima piña (Venezuela); cascabel púa, pucarara, matacaballo (Bolivia); surucucu, surucutinga, pico-de-jaca, surucucu-pico-de-jaca (Brazil).

Recognition: ♂♂ 340 cmMaximum distance from the snout to the tip of the tail. ♀♀ 291 cmMaximum distance from the snout to the tip of the tail.. The Amazonian Bushmaster (Lachesis muta) is probably the longest venomous snake in the Americas, and the second in the world, after the King Cobra (Ophiophagus hannah).^1,2 The great majority of adults of L. muta measure less than 2.5 m, but some individuals measuring up to 4.5 m have been reported, but not confirmed.³ In the Ecuadorian Amazon, L. muta is identified from other vipers by having the following features: tail ending in a spine, prominent knoblike keels on the dorsal scales, which give the snake a “pineapple texture,” and a distinctive color pattern consisting of bold-black inverted triangles on a light brown ground dorsum (Fig. 1).^2,4,5 Neonates (40–54.2 cm in total length)^1,6 have a vibrant orangish dorsum and a yellowish tail.² The most similar viper species that may be found living alongside L. muta in Ecuador is Bothrocophias microphthalmus, which is distinguished by having a dorsal pattern consisting of X-shaped markings on a brownish background.

Figure 1: Individuals of Lachesis muta from Llanganates National Park, Pastaza province, Ecuador.

Natural history: RareTotal average number of reported observations per locality less than ten. to extremely rareTotal average number of reported observations per locality less than three.. Lachesis muta is a nocturnal, solitary, and terrestrial viper that inhabits old-growth to moderately disturbed evergreen lowland and foothill forests, tropical dry forests and savannas (in Brazil),⁷ swamps,² and plantations (of bamboo, banana, cacao, palm-oil, rubber, sugarcane, tea, or yucca).^8–12 Occasionally, Amazonian Bushmasters venture into pastures,⁹ rural gardens, and houses.^13,14 This species seems to prefer upland forest areas, but may as well occur in seasonally flooded forests during low-water season.^2,15

Adult individuals of Lachesis muta spend most of their time (during the day as well as at night) coiled in ambush posture on the forest floor, either exposed or under herbaceous vegetation.^13,16 They are also occasionally seen crawling on the forest floor,² swimming across rivers,¹⁷ or crossing roads in the vicinity of forested areas.^13,16 Juveniles dwell on leaf-litter, logs, buttress roots, and rocky areas.⁶ When not active, Amazonian Bushmasters hide under big rocks,¹⁸ under logs,¹⁶ in spiny palm scrub,¹⁹ or in any hole that fits the size and does not flood,¹⁸ particularly caves by armadillos, agoutis, and rabbits.^13,18,20 Individuals may remain hidden in these lairs for days or even weeks.²

Individuals of Lachesis muta rely on their camouflage as a primary defense mechanism. Unlike popular belief, they are not aggressive, but calm and sluggish when confronted by humans.^9,14 In the presence of a disturbance, most individuals show no immediate defensive reaction, remaining coiled and immobile even when the vegetation around the snake is being cleared.¹³ If disturbed further, most Amazonian Bushmasters will flee, while others might emit a low whistle-like sound, vigorously wiggle the tail against the leaf-litter, or readily strike without a warning.⁸

In Ecuador and Peru, some indigenous groups (Achuar, Kichwa, Sápara, Shiwiar, and Shuar) associate the call of two species of Amazonian treefrogs (Tepuihyla tuberculosa and T. shushupe), with the “calling” of the Amazonian Bushmaster.^9,21 The sound can be described as “an unearthly ascending series of kaks.”² The origin of this association is unsure, since individuals of Lachesis muta cannot produce loud sounds.

South American Bushmasters are ambush predators. They sit-and-wait for prey to pass by, usually along small mammal pathways. Depending on the relative size of the prey, the snakes can “bite and hold” their prey or “bite and release,” subsequently following the scent trail of the envenomated prey.²² The diet of Lachesis muta is primarily based on small and medium-sized mammals such as rodents (rice rats, spiny rats, and agoutis),^16,23,24 porcupines, squirrels, and opossums,^2,25 but it also occasionally includes squirrel monkeys, frogs, and birds.^26,27

Predators of Lachesis muta include snakes (Clelia clelia),²⁸ peccaries,²⁹ and humans.² Indigenous communities in the Amazon consider Lachesis a delicacy, and there are records of bushmasters being cooked in villages in Peru² and Venezuela.³⁰ Individuals are often parasitized by tongue worms (Pentastomida) and lung worms (Rhabditida), which impairs respiration and may cause infected individuals to die from pneumonia.¹

Lachesis muta is a venomous species (LD50The median lethal dose (LD50) is a measure of venom strength. It is the minium dosage of venom that will lead to the deaths of 50% of the tested population. 2.8–9.75 mg/kg)^31–33 that is responsible for 4.5–7.5% of snakebites throughout its range.^2,14,34,35 In some areas, 17–35% of snakebites are attributed to this species, but these numbers almost surely correspond to misidentifications or exaggerations.^14,36 Human envenomations by L. muta, although infrequent,³⁶ can be severe due to the high amount of venom inoculated (up to 552 mg of dry venom per bite).³⁷

The venom of Lachesis is noteworthy because in addition to being hemorrhagic, necrotizing, myotoxic, and proteolytic, is also neurotoxic.^36,39 Human victims of L. muta snakebite may experience agonizing local pain, swelling, bleeding, blistering, incoagulable blood, and necrosis (death of tissues and cells).^14,31,40 Most victims also experience a syndrome usually not seen in those bitten by other viper species. This so-called “Lachesis Syndrome” includes nausea, vomiting, diarrhea, sweating, low blood pressure, impaired consciousness, faintness, respiratory distress, and shock.³⁶

In poorly managed or untreated cases, envenomations by Lachesis muta can result in secondary infection, septicemia, shock, pulmonary edema, renal failure, amputations, permanent crippling deformity, disabilities, and (in 1.4% of cases)⁴¹ death in as little as 45 minutes.^2,14,34,42 However, some bites to humans involve no envenomation at all (“dry bites”).⁴³

What to do if you are bitten by a Amazonian Bushmaster? Remain calm. Remove rings and tight fitting clothes to avoid swelling. Reduce movement of bitten extremity to reduce absorption of venom. Avoid the application of tourniquets, electric shocks, traditional medicine, venom suction, and incision of the bite wound. Plan immediate evacuation to a medical facility that has antivenom and avoid any action that may delay transportation. At the medical facility, personnel can initiate treatment with the appropriate antivenom, monitor vital signs, and perform laboratory tests.

Fortunately, the antivenom that is available in Ecuador, although produced based on individuals from other countries, can neutralize the venom of Lachesis muta.^35,44 This is because the venom’s toxic and enzymatic activities are similar across Lachesis populations.⁴⁴ Although serum therapy (antivenom) is the only recommended approach against a bite by an Amazonian Bushmaster, the fruit juice and the aqueous extract of leaves of soursop (Annona muricata) used by traditional “healers” may exert a protective action against the myotoxicity (muscle-breaking activity) of the venom.⁴⁵

Breeding in Lachesis muta appears to be triggered by cold fronts and storms rather than correspond to a specific time of the year.^6,46 During these times, males actively follow the pheromone trail of females.^4,6 They also vigorously fight other males over access to females.¹ During combats, males entwine their bodies with each other and try to push the opponent against the substrate.¹ Courtship can last up to five hours and consists of the male rubbing its head and body, while flicking the tongue, over the female's body.⁴ Bushmasters are unique among New World vipers in that they lay eggs rather than give birth to live young.^2,47 Females are capable of delaying fertilization by storing sperm for months.⁴⁸ They deposit 3–23 eggs^15,49,50 measuring 7–10 cm long⁵⁰ in hollow logs or in mammal burrows.^2,20

Unlike other South American vipers, females of Lachesis muta exhibit maternal care. They protect their clutch during the entirety of the incubation period, which is 60–95 days (~2–3 months).^48,50 Considering the gestation plus incubation period, females can spend up to seven months without feeding.⁴⁸ Hatchlings measure 40–54.2 cm in total length.^1,6 Males become sexually mature when they reach ~122–129 cm in total length; females at ~112–164 cm, or in about 1–2 years.^48,50,51 Under human care, individuals can live up to ~16 years,² and probably much longer.

Conservation: Least Concern Believed to be safe from extinction given current circumstances.. Lachesis muta is here proposed to be listed in this category because it is widely distributed,⁷ especially throughout the Amazon basin, a region that retains most of its original forest cover. Therefore, we consider that the species faces no major immediate extinction threats. We estimate that, in Ecuador, ~87.3% of the habitat of L. muta holds pristine forest. Unfortunately, the population of L. muta occurring on the Atlantic Forest of Brazil, which is genetically distinct,⁴⁴ occurs over an area that has lost the majority (84–88.6%) of its forest cover,⁵² and is now largely restricted to forest fragments.^53,54 Habitat destruction and consequent habitat fragmentation is the main threat to the long-term survival of the species.⁵⁴ Other threats include traffic mortality,^1,55,56 decline in small mammal populations, and direct killing.⁵⁴

Distribution: Lachesis muta is native to an estimated ~5,489,350 km² area throughout the Amazon basin and adjacent foothills of the Andes in Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, and Venezuela.⁷ It also occurs in the Atlantic Forest of Brazil.⁷ In Ecuador, the species occurs at elevations between 106 and 1568 m (Fig. 2).

Figure 2: Distribution of Lachesis muta in Ecuador. See Appendix 1 for a complete list of the presence localities included in the map.

Etymology: The generic name Lachesis is derived from the name of one of the “Three Fates” (also known as Moirae or incarnations of destiny) in Greek mythology.² Lachesis was the Moira that decided the length of life allotted to each human being and god.² This epithet was probably given to reflect the feeling of having one’s fate at the mercy of the snake upon encountering such an imposing creature.⁵⁷ The specific epithet muta is derived from the Latin word mutus (meaning “dumb” or “silent”).⁵⁸ It refers the lack of a rattle, which is present in snakes of the closely related snake genus Crotalus.²

See it in the wild: In Ecuador, Amazonian Bushmasters are recorded rarely, usually no more than once every few months at any given locality. However, in areas having unspoiled forest and low human population density, like in Yasuní Scientific Station and Bigal River Biological Reserve, individuals may be seen more frequently, albeit certainly not reliably. The snakes may be located by walking along trails or by cruising roads through primary forest.

FREQUENTLY ASKED QUESTIONS:

How long is a bushmaster snake? Amazonian Bushmasters can grow up 3.4 m, which probably makes them the longest vipers in the world, and the second longest venomous snakes in the world, after the King Cobra (Ophiophagus hannah).1,2

How venomous is the Amazonian Bushmaster? With a lethal dose of LD50The median lethal dose (LD50) is a measure of venom strength. It is the minium dosage of venom that will lead to the deaths of 50% of the tested population. 2.8–9.75 mg/kg,31–33 the venom of the Amazonian Bushmaster is considered “extremely toxic.” In poorly managed or untreated human envenomations, the venom may cause renal failure, amputations, permanent crippling deformity, disabilities, and (in 1.4% of cases)41 death in as little as 45 minutes.2,14,34,42

Where are bushmasters snakes found? Bushmaster snakes occur in the tropical forests of Central American and northern South America.2

What does the Amazonian Bushmaster eat? Amazonian Bushmasters feed primarily on small and medium-sized mammals such as rodents (rice-rats, spiny rats, and agoutis),16,23,24 porcupines, squirrels, and opossums,2,25 but they also occasionally include squirrel monkeys, frogs, and birds in their diet.26,27

Will a bushmaster snake chase you? There are records of bushmaters actively, albeit slowly, pursuing people,1,5,38 but these instances probably correspond to snakes that mistake a warm foot or warm hand for food.2

Acknowledgments: The first author would like to thank Caroline Guevara-Molina for the comments and suggestions that enhanced the scientific quality of this work. Special thanks to Andreas Kay, Antonio Freire, Darwin Núñez, David Salazar, and Regdy Vera, for providing locality data for Lachesis muta. For providing natural history data, we are particularly grateful to Darwin Núñez, Ernesto Arbeláez, Harry Turner, and Konrad Mebert. The second author would like to thank Christopher Gillette for providing an image of an Amazonian Bushmaster preying upon a rice rat.

Authors: Juan C. Díaz-Ricaurte,^aAffiliation: Universidade de São Paulo, São Paulo, Brazil. Alejandro Arteaga,^bAffiliation: Tropical Herping (TH), Quito, Ecuador. and Juan M Guayasamin^cAffiliation: Laboratorio de Biología Evolutiva, Universidad San Francisco de Quito (USFQ), Quito, Ecuador.^,dAffiliation: Galapagos Science Center, Galápagos, Ecuador.^,eAffiliation: Centro de Investigación de la Biodiversidad y Cambio Climático, Universidad Tecnológica Indoamérica, Quito, Ecuador.

Photographers: Jose Vieira^bAffiliation: Tropical Herping (TH), Quito, Ecuador.^,fAffiliation: ExSitu, Quito, Ecuador.

How to cite? Díaz-Ricaurte JC, Arteaga A, Guayasamin JM (2020) Amazonian Bushmaster (Lachesis muta). In: Arteaga A, Bustamante L, Vieira J (Eds) Reptiles of Ecuador: Life in the middle of the world. Available from: www.reptilesofecuador.com. DOI: 10.47051/BJCI8462

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Appendix 1: Locality data used to create the distribution map of Lachesis muta in Ecuador (Fig. 2). Go to the section on symbols and abbreviations for a list of acronyms used.

Country	Province	Locality	Source
Colombia	Caquetá	Bocana Camicaya	Díaz-Ricaurte et al. 2017
Colombia	Caquetá	La Floresta	Díaz-Ricaurte et al. 2017
Colombia	Caquetá	La Paz	Díaz-Ricaurte et al. 2017
Colombia	Caquetá	Paujiles	Díaz-Ricaurte et al. 2017
Colombia	Caquetá	Pore	Díaz-Ricaurte et al. 2017
Colombia	Caquetá	San Isidro	Díaz-Ricaurte et al. 2017
Colombia	Caquetá	Santa Fe	Díaz-Ricaurte et al. 2017
Colombia	Putumayo	Bloque 24	Chávez 2016
Colombia	Putumayo	La Hormiga	Díaz-Ricaurte et al. 2017
Ecuador	Morona Santiago	Below Alshi	MAE Morona Santiago
Ecuador	Morona Santiago	Bomboiza	Terán & Lomonte 2016
Ecuador	Morona Santiago	Centro Chuwints	Nogueira et al. 2019
Ecuador	Morona Santiago	Centro Shuar Kenkuim	Valencia et al. 2016
Ecuador	Morona Santiago	Centro Shuar Kiim	Valencia et al. 2016
Ecuador	Morona Santiago	Chuwints	Valencia et al. 2016
Ecuador	Morona Santiago	Cueva de los Tayos	DHMECN 64
Ecuador	Morona Santiago	Cusuime	Ortega-Andrade 2010
Ecuador	Morona Santiago	El Quimi	Betancourt et al. 2018
Ecuador	Morona Santiago	El Tiink	This work
Ecuador	Morona Santiago	Fauna de la Amazonía	Andreas Kay, pers. comm.
Ecuador	Morona Santiago	Gualaquiza	Antonio Freire, pers. comm.
Ecuador	Morona Santiago	Kankaim	Valencia et al. 2016
Ecuador	Morona Santiago	Limón	Nogueira et al. 2019
Ecuador	Morona Santiago	Makuma	Campbell & Lamar 2004
Ecuador	Morona Santiago	Mutintz	Valencia et al. 2016
Ecuador	Morona Santiago	Nunkandai	Valencia et al. 2016
Ecuador	Morona Santiago	Quebrada Río Napinaza	David Salazar, pers. comm.
Ecuador	Morona Santiago	Reserva el Quimi	iNaturalist
Ecuador	Morona Santiago	San Carlos de Limón	Valencia et al. 2016
Ecuador	Morona Santiago	San Karlos	Peñafiel 2013
Ecuador	Morona Santiago	Tunants	Valencia et al. 2016
Ecuador	Morona Santiago	Wisui	Chaparro et al. 2011
Ecuador	Morona Santiago	Yukaip	Nogueira et al. 2019
Ecuador	Napo	Huaorani Lodge	Photo record by Peter Horrell
Ecuador	Napo	Huataraco	Valencia et al. 2016
Ecuador	Napo	Jatun Sacha	Vigle 2008
Ecuador	Napo	Río Pusuno	iNaturalist
Ecuador	Napo	San Jose de Dahuano	Valencia et al. 2016
Ecuador	Napo	Shiguancocha	Photo record by Fausto Gómez
Ecuador	Napo	Wild Sumaco	Camper et al. (in press)
Ecuador	Orellana	130 km S Tiguino	USNM 321138
Ecuador	Orellana	Bloque 31	Libro PetroAmazonas
Ecuador	Orellana	Bodega NPW	This work
Ecuador	Orellana	Coca	MHNG 2410.043
Ecuador	Orellana	Cotapino	Nogueira et al. 2019
Ecuador	Orellana	Loreto	Nogueira et al. 2019
Ecuador	Orellana	NPF–Tivacuno, km 7 1/2	Nogueira et al. 2019
Ecuador	Orellana	Parcela 50 ha	Nogueira et al. 2019
Ecuador	Orellana	Pompeya Sur	Nogueira et al. 2019
Ecuador	Orellana	Río Bigal	Photo record by Thierry García
Ecuador	Orellana	San Jose de Mote	iNaturalist
Ecuador	Orellana	Shiripuno Lodge	Photo record by Fernando Vaca
Ecuador	Orellana	Tambococha	Nogueira et al. 2019
Ecuador	Orellana	Tiputini Biodiversity Station	Nogueira et al. 2019
Ecuador	Orellana	Vía Pompeya Sur-Iro, km 58	Nogueira et al. 2019
Ecuador	Orellana	W Puerto Ventura	iNaturalist
Ecuador	Orellana	Waimo of Conoco	Zamudio & Greene 1997
Ecuador	Pastaza	Balsaura	Ortega-Andrade 2010
Ecuador	Pastaza	Conambo	Ortega-Andrade 2010
Ecuador	Pastaza	Copataza (Achuar)	Peñafiel 2013
Ecuador	Pastaza	Intersección Cueva de los Tayos	Peñafiel 2013
Ecuador	Pastaza	Juyuintza	Ortega-Andrade 2010
Ecuador	Pastaza	Kapawi Lodge	This work
Ecuador	Pastaza	Kurintza	Ortega-Andrade 2010
Ecuador	Pastaza	Latasas–Umupi	Morillo-Tobar 2013
Ecuador	Pastaza	Paparawua	Nogueira et al. 2019
Ecuador	Pastaza	Peter Archer's place	Photo record by Peter Archer
Ecuador	Pastaza	Piatúa	This work
Ecuador	Pastaza	Pindoyacu	Ortega-Andrade 2010
Ecuador	Pastaza	Río Bufeo	Ortega-Andrade 2010
Ecuador	Pastaza	Río Chambira	AMNH 49135
Ecuador	Pastaza	Río Pucayacu	USNM 165010
Ecuador	Pastaza	Río Villano	AMNH 82006
Ecuador	Pastaza	Sumak Kawsay	Photo record by Vincent Premel
Ecuador	Pastaza	Tzarentza	This work
Ecuador	Sucumbíos	Baboroé	Yánez-Muñoz & Chimbo 2007
Ecuador	Sucumbíos	Batallón de Selva 56	Photo record by Guido Bladimir
Ecuador	Sucumbíos	Estación PUCE Cuyabeno	Nogueira et al. 2019
Ecuador	Sucumbíos	La Selva Lodge	Nogueira et al. 2019
Ecuador	Sucumbíos	Lago Agrio	Nogueira et al. 2019
Ecuador	Sucumbíos	Limoncocha	UF 30615
Ecuador	Sucumbíos	Napo Wildlife Center	This work
Ecuador	Sucumbíos	Pañacocha	Nogueira et al. 2019
Ecuador	Sucumbíos	Pisorié	Yánez-Muñoz & Chimbo 2007
Ecuador	Sucumbíos	Reserva Cuyabeno	Nogueira et al. 2019
Ecuador	Sucumbíos	Sacha Lodge	Nogueira et al. 2019
Ecuador	Sucumbíos	San Pablo de Kantesiya	MECN 65
Ecuador	Sucumbíos	Sani Lodge	This work
Ecuador	Sucumbíos	Santa Cecilia	Duellman 1978
Ecuador	Sucumbíos	Shishicho	Pitman et al. 2002
Ecuador	Zamora Chinchipe	Bombuscaro	Darwin Núñez, pers. comm.
Ecuador	Zamora Chinchipe	Los Encuentros	Darwin Núñez, pers. comm.
Ecuador	Zamora Chinchipe	Parque Nacional Podocarpus 1	iNaturalist
Ecuador	Zamora Chinchipe	Parque Nacional Podocarpus 2	iNaturalist
Ecuador	Zamora Chinchipe	Pindal–Machinaza	MZUTI 5501
Ecuador	Zamora Chinchipe	Sendero Ciudad Perdida	Darwin Núñez, pers. comm.
Ecuador	Zamora Chinchipe	Tundayme	This work
Ecuador	Zamora Chinchipe	Zamora	Darwin Núñez, pers. comm.
Perú	Amazonas	Alto Cenepa	iNaturalist
Perú	Amazonas	Chosica	Consorcio AGUA SELVA 2014
Perú	Amazonas	Cordillera del Cóndor	RAP 1994
Perú	Loreto	Alto Cahuapanas	Venegas et al. 2014
Perú	Loreto	N of Charupa	Campbell & Lamar 2004
Perú	Loreto	Pavayacu	Campbell & Lamar 2004
Perú	Loreto	Pongo Chinim	FMNH 2012
Perú	Loreto	Ucayali	iNaturalist
Perú	Loreto	Yarina Cocha	Campbell & Lamar 2004

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