Bothriopsis bilineataGreen Jararaca

Ge­o­graphic Range

Both­ri­op­sis bi­lin­eata is found in the South Amer­i­can neotrop­ics. The largest pop­u­la­tions are found in the Ama­zon­ian re­gions of Bo­livia, Co­lum­bia, Equador, Peru, and Brazil. Other highly pop­u­lated re­gions in­clude equa­to­r­ial forests of Venezuela, Guyana, French Guiana, and Suri­name. It is most abun­dant in south­west South Amer­ica, in Par­que Na­cional. Both­ri­op­sis bi­lin­eata has been doc­u­mented as far south as Jacarepagua, which is lo­cated within the city lim­its of Rio de Janeiro. How­ever, the species was re­moved from Jacarepagua in 1963 and is now con­sid­ered rare in that re­gion. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004)

Habi­tat

Two-striped for­est-pitvipers are pre­sent only in forested habi­tats. Its dis­tri­b­u­tion ranges from sea level to ap­prox­i­mately 1000 m in el­e­va­tion. They very sel­dom are found at el­e­va­tions above 1000 m. Their most com­mon habi­tat is wet low­land rain­forests, where they live pri­mar­ily near wa­ter­ways, es­pe­cially along streams. Two-striped for­est-pitvipers spend much of their time on branches and shrubs more than 1 m off the ground. While they are par­tic­u­larly abun­dant in very wet pri­mary forests, these snakes also occur in sec­ondary forests near or ad­ja­cent to pri­mary forests. Wet low­land rain­forests oc­cu­pied by two-striped for­est-pitvipers usu­ally have a canopy height of 40 m, but in­di­vid­ual trees can reach up to 65 m. How­ever, trees in wet low­land rain­forests typ­i­cally have a greater height, height to first branch, trunk vol­ume, but­tress height and but­tress area than in other rain­for­est ecosys­tems. Sub-canopy veg­e­ta­tion in low­land rain­forests pri­mar­ily con­sists of suc­ces­sional trees rather than shrubs or herbs. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Daly and Mith­cell, 2000)

  • Range elevation
    0 to 1000 m
    0.00 to 3280.84 ft

Phys­i­cal De­scrip­tion

Both­ri­op­sis bi­lin­eata is a sleek, green pitviper with a pre­hen­sile tail. Typ­i­cal length is about 70 cm, al­though in­di­vid­u­als up to 123 cm have been re­ported. Fe­males are usu­ally 10 to 20 cm longer than males. In both sexes, the head is typ­i­cally green with black spots, but some­times may be green with tan or red spots with black mar­gins. The iris of the eye is light green to yel­low and the pupil is ver­ti­cally el­lip­ti­cal. The tongue is pink­ish-grey with dark grey tips. Snout length in B. bi­lin­eata is sim­i­lar to that of other pit vipers, with a well de­fined can­thus ros­tralis. Body col­oration of in B. bi­lin­eata is pale green with black, red, or tan spots, on the dor­sum. The en­larged scales on ven­ter, com­monly re­ferred to as ven­trals, are yel­low with a green tinged bor­der­ing. A creamy yel­low line bor­ders the lat­eral mar­gins of the ven­trals and runs the en­tire length of the snake. The dis­tal por­tion of the tail is pink out­lined by yel­low. In­di­vid­u­als with a whitish-yel­low tail tip have been re­ported. (Mon­teiro, et al., 2006; Val­dujo, et al., 2002; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Mon­teiro, et al., 2006; Val­dujo, et al., 2002)

The two-striped for­est pitviper are pri­mar­ily ar­bo­real. They weigh less and are a no­tably shorter in length than other species of pitviper. This dif­fer­ence in length has been at­trib­uted both to their ar­bo­real habi­tat and feed­ing be­hav­ior. Smaller body size al­lows them to ar­tic­u­late through the dense shrub­bery of the Ama­zon re­gion. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Mon­teiro, et al., 2006; Val­dujo, et al., 2002)

Other char­ac­ter­is­tic fea­tures of Both­ri­op­sis bi­lin­eata in­clude 5 to 9 keeled in­ter­supraoc­u­lars, 7 to 9 supral­abi­als, 8 to 12 in­fral­abi­als, and 23 to 25 mid­body scale rows. Males have 190 to 218 ven­trals and 65 to 76 mostly di­vided sub­cau­dals, whereas fe­males have 192 to 220 ven­trals and 55 to 73 mostly di­vided sub­cau­dals. Usu­ally B. bi­lin­eata has a la­cuno­labial that is formed by the sec­ond supral­abial and prela­cu­nal fused to­gether. How­ever, some­times pre­sent are par­tial or full su­tures that sep­a­rate these scales form­ing the la­cuno­labial. Mem­bers of the genus Both­ri­op­sis have unique skull mor­phol­ogy in that the frontal bones are con­cave and are longer than than they are wide. The post­frontals do not con­tact the frontal bones and are long and nar­row. All pitvipers have 5 pala­tine teeth, 14 to 18 ptery­goid teeth, and 15 to 17 den­tary teeth. Their max­il­lary fangs are around 1.5 to 2 times longer than the height of the max­il­lary bone. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Mon­teiro, et al., 2006; Val­dujo, et al., 2002)

Re­lated or sim­i­lar species to Both­ri­op­sis bi­lin­eata in­cludes sub­species B. b. smaragdi­nus, which dif­fers in that it does not pos­sess tan or red spots on its body. Rather, it is pep­pered only with black spots. One other pre­hen­sile-tailed pitviper in South Amer­ica that might be con­fused with B. bi­lin­eata is Both­riechis schlegelii, which has white or yel­low lines. How­ever, sym­pa­try be­tween Both­ri­op­sis bi­lin­eata and Both­riechis schlegelii has not been con­firmed. In Ecuador, Peru, and Bo­livia, there are four other green pitvipers. These in­clude Both­ri­op­sis pul­chra, B. pul­chra, B. pe­ru­viana, and B. oligolep­sis. All of these species lack the par­aven­tral stripes found in B. bi­lin­eata. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Mon­teiro, et al., 2006; Val­dujo, et al., 2002)

  • Sexual Dimorphism
  • female larger
  • Range length
    57.3 to 123 cm
    22.56 to 48.43 in
  • Average length
    <70cm cm
    in

De­vel­op­ment

Both­ri­op­sis bi­lin­eata is ovo­vi­vip­a­rous, mean­ing that the eggs de­velop in­ter­nally, with the mother giv­ing birth to live young. Neonates are ven­omous upon birth and are known to be more ag­gres­sive at times than the adults. After birth, young feed upon frogs, lizards, and other rel­a­tively small prey. In­di­vid­ual size de­pends greatly on food avail­abil­ity. Adult fe­males are typ­i­cally larger than adult males. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Tra­vers, 1999)

Re­pro­duc­tion

There is no in­for­ma­tion avail­able re­gard­ing the mat­ing sys­tem of Both­ri­op­sis bi­lin­eata. How­ever, a study of Both­ri­op­sis tae­ni­ata at the Dal­las Zoo found that, dur­ing mat­ing, males po­si­tion them­selves on one side of the fe­male, while mov­ing across her dor­sum. While mov­ing, the male makes very jerky mo­tions and a hemipe­nis into the fe­male’s cloaca. The male flicks his tongue ex­tremely fast, ap­prox­i­mately 43 to 71 times a minute. Dur­ing this study, fe­males gave birth in June, July, and Oc­to­ber, with clutches rang­ing in size from 7 to 12 off­spring. In many pitvipers, male-male com­pe­ti­tion, known as "the dance of the adders", takes places dur­ing the mat­ing sea­son and is mainly pre­formed to guard ac­cess to mates. Some re­ports sug­gest that this be­hav­ior may also be used to com­pete for ter­ri­to­ries. Fight­ing be­tween males is done by hold­ing the face and fore­body erect and wrap­ping their bod­ies around one an­other as they at­tempt to force one an­other to the ground. Com­pet­i­tive bouts be­tween males can take hours and usu­ally ends with the win­ner gain­ing ac­cess a nearby fe­male for mat­ing. (Al­me­dia-San­tos and Sa­lo­mao, 2002; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Tra­vers, 1999)

Both­ri­op­sis bi­lin­eata is ovo­vi­vip­a­rous, mean­ing that the eggs de­velop in­ter­nally, with the mother giv­ing birth to live young. Clutch size ranges from 6 to 16 young. Neonates range in length from 20 to 26 cm and range in mass from 7 to 11 g, with an av­er­age of 10 g. Al­though lit­tle in­for­ma­tion is known con­cern­ing the re­pro­duc­tive be­hav­ior of Both­ri­op­sis bi­lin­eata, re­pro­duc­tive be­hav­ior of north­ern tem­per­ate pitvipers is thought to be very sim­i­lar. Re­pro­duc­tive cycle be­gins with sper­mato­ge­n­e­sis and vitel­lo­ge­n­e­sis. Vitel­lo­ge­n­e­sis in fe­males usu­ally oc­curs dur­ing late sum­mer or fall with the de­vel­op­ment of fol­li­cles paus­ing dur­ing win­ter and re­sum­ing dur­ing spring. Courtship and mat­ing gen­er­ally oc­curs dur­ing late sum­mer to win­ter, from April to Oc­to­ber. Long-term sperm stor­age may be used dur­ing the win­ter, from July to Sep­tem­ber, and is es­sen­tial in many pitvipers. De­lay­ing fer­til­iza­tion makes it pos­si­ble for birthing to occur dur­ing a fa­vor­able time of year and likely in­creases off­spring sur­vival. Dur­ing spring, ovu­la­tion, fer­til­iza­tion and em­bryo­ge­n­e­sis take place. Birth most likely oc­curs dur­ing the rainy sea­son of sum­mer, which is usu­ally in March. How­ever, this cycle likely varies in re­la­tion to weather, food avail­abil­ity and by species. (Al­me­dia-San­tos and Sa­lo­mao, 2002; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Tra­vers, 1999)

  • Range number of offspring
    4 to 16
  • Average number of offspring
    9

Both­ri­op­sis bi­lin­eata fe­males store sperm until the op­ti­mal time to fer­til­ize the eggs. Once the eggs are fer­til­ized, the mother car­ries the em­bryos until the eggs are ready to hatch, which oc­curs in­ter­nally. For all pitvipers, young ven­omous upon birth and are able to hunt. Some moth­ers stay with the young for 7 to 10 days. How­ever, young are typ­i­cally in­de­pen­dent upon birth. (Al­me­dia-San­tos and Sa­lo­mao, 2002; Tra­vers, 1999)

  • Parental Investment
  • pre-hatching/birth
    • provisioning
      • female

Lifes­pan/Longevity

There is no in­for­ma­tion on the av­er­age lifes­pan of Both­ri­op­sis bi­lin­eata. In gen­eral, es­ti­mated lifes­pan for pitvipers ranges from 10 to 25 years, with vari­a­tions most likely due to cap­tiv­ity. One ac­count de­scribes a B. tae­ni­ata that lived in the Dal­las Zoo for 13 years, after being re­ceived by the zoo as an adult. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Tra­vers, 1999)

Be­hav­ior

Both­ri­op­sis bi­lin­eata is noc­tur­nal and spends most of the day in thick fo­liage or at the base of palm trees, gen­er­ally 1 m or higher above the ground. It uses its pre­hen­sile tail to an­chor it­self on to trees and shrub­bery. To con­serve en­ergy, B. bi­lin­eata tends to hide in shaded areas and moves very lit­tle through­out the day. Both­ri­op­sis bi­lin­eata is an am­bush preda­tor. Like other pitvipers, B. bi­lin­eata has pits re­sem­bling nos­trils that sense var­i­ous wave­lengths of in­frared heat. Be­cause tem­per­a­tures at night are cooler, a greater heat sig­na­ture is gen­er­ated by po­ten­tial prey. The pits are lo­cated on both sides of the head, cre­at­ing a stereo ef­fect that al­lows B. bi­lin­eata to de­ter­mine the di­rec­tion and dis­tance of prey. Ex­per­i­ments con­ducted on pitvipers have shown that when de­prived of sight and smell, they can strike ac­cu­rately on tar­gets 0.2 C greater than sur­round­ing am­bi­ent tem­per­a­ture. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Tra­vers, 1999)

Home Range

There is no in­for­ma­tion avail­able re­gard­ing the av­er­age home range size of Both­ri­op­sis bi­lin­eata.

Com­mu­ni­ca­tion and Per­cep­tion

Both­ri­op­sis bi­lin­eata, like other pitvipers, has heat-sens­ing pits lo­cated be­tween the eyes and nos­trils, on both sides of the head. Within these pits are nu­mer­ous heat-sen­si­tive nerves that are cov­ered by thin mem­bra­nous tis­sue. The pits are highly vas­cu­lar­ized, which helps main­tain a con­stant tem­per­a­ture, thereby in­creas­ing sen­si­tiv­ity to changes in am­bi­ent tem­per­a­ture. In ad­di­tion to heat-sens­ing pits, B. bi­lin­eata also has well-de­vel­oped binoc­u­lar vi­sion re­sult­ing from its ver­ti­cal slits. This trait, com­mon to all pitvipers, in­creases depth and vi­sual per­cep­tion. Like all snakes, Both­ri­op­sis bi­lin­eata has an adept ol­fac­tory per­cep­tion. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004)

Other than those that occur dur­ing mat­ing sea­son, lit­tle is known about in­traspe­cific in­ter­ac­tions in Both­ri­op­sis bi­lin­eata. Dur­ing mat­ing sea­son, male-male com­pe­ti­tion takes place. Dur­ing these com­pet­i­tive in­ter­ac­tions, two males be­come in­ter­twined in an at­tempt to force one an­other to the ground. The win­ner gains ac­cess to mates and ter­ri­tory. Al­though they lack ears, snakes are highly sen­si­tive to vi­bra­tions. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004)

Food Habits

Two-striped for­est pitvipers are noc­tur­nal preda­tors. They rest in trees and shrubs 3 to 4 feet above the ground and use their heat-sens­ing pits to gen­er­ate a ther­mal pic­ture of the sur­round­ing en­vi­ron­ment and po­ten­tial prey. Ev­i­dence sug­gests that they oc­ca­sion­ally use their tails as a lure to at­tract prey. These vipers prey mainly on frogs, but also reg­u­larly eat small birds. Re­cently, stom­ach con­tent analy­sis has re­vealed that two-striped for­est pitvipers also feed on a va­ri­ety of lizards and small mam­mals, in­clud­ing mice and rab­bits. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Ro­drigo C. G. de Souza, M.D. and Fun­daçã o Hos­pi­ta­lar Itacare, 2007)

  • Primary Diet
  • carnivore
    • eats terrestrial vertebrates
  • Animal Foods
  • birds
  • mammals
  • amphibians
  • reptiles

Pre­da­tion

Neotrop­i­cal birds of prey tend to feed on ven­omous snakes, but be­cause two-striped for­est pitvipers are noc­tur­nal and well-cam­ou­flaged, lit­tle is known about their preda­tors. Birds of prey known to feed on ven­omous snakes in the Ama­zon in­clude laugh­ing fal­cons, crane hawks, road­side hawks, and swal­low-tailed kites. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; "De­for­esta­tion in the Ama­zon", 2010)

  • Anti-predator Adaptations
  • cryptic

Ecosys­tem Roles

Both­ri­op­sis bi­lin­eata are preda­tors of many species of small birds, frogs, and lizards in their ecosys­tem. They are highly ven­omous and are prob­a­bly avoided by most species through­out their ge­o­graphic range. Par­a­sites of this species are un­known. (Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

In the past, the venom of Both­ri­op­sis bi­lin­eata has been the sub­ject of re­search for its an­ti­co­ag­u­la­tion prop­er­ties; how­ever, lo­cat­ing this species is re­port­edly very chal­leng­ing. There­fore, not a great deal of re­search has been con­ducted on this species. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004)

  • Positive Impacts
  • research and education

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Both­ri­op­sis bi­lin­eata is po­ten­tially dan­ger­ous, as a sin­gle strike from this snake re­leases ex­tremely toxic venom. Their venom is con­tains pro-co­ag­u­lant and an­ti­co­ag­u­lant agents and their bite can re­sult in se­vere he­m­or­rhag­ing. Both­ri­op­sis bi­lin­eata has been re­ported to be the sec­ond most dan­ger­ous snake in the Ama­zon, be­hind only Both­rops atrox. Nu­mer­ous ac­counts of bites have been doc­u­mented, some re­sult­ing in death. In stud­ies of snake bite cases near Co­lum­bia, about 15% of bites were from B. bi­lin­eata. Bleed­ing from the gums, site of bite, and ori­fices often occur after a bite. Other re­lated symp­toms in­clude he­mateme­sis, hema­turia, loss of con­scious­ness, ery­thema, headaches, pain at en­ven­o­ma­tion site, tis­sue necro­sis, and edema. Both­ri­op­sis bi­lin­eata is not con­sid­ered an ag­gres­sive species, but has been known to strike when dis­turbed. Be­cause this is an ar­bo­real species, most bites re­ported are to the hands, arms, and face. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; O'Shea, 2005; Small­i­gan, et al., 2004; Tra­vers, 1999)

  • Negative Impacts
  • injures humans

Con­ser­va­tion Sta­tus

Both­ri­op­sis bi­lin­eata has not been eval­u­ated by the IUCN and is not cur­rently listed on any threat­ened species lists. The great­est threat to this species is hu­mans. De­for­esta­tion has been linked to pop­u­la­tion de­clines, and from 1988 to 2010, about 385,000 km^2 of pri­mary habi­tat in the Ama­zon has been de­stroyed. While it is know that this species is dis­trib­uted through­out low­land rain forests in the Ama­zon, Bo­livia, Peru, Ecuador, Brazil, Venezuela, and the Guianas (el­e­va­tions below 1,000 m, usu­ally near wa­ter­ways), lit­tle in­for­ma­tion is avail­able about cur­rent pop­u­la­tion sizes. Its noc­tur­nal ac­tiv­ity and col­oration make its num­bers dif­fi­cult to es­ti­mate. Be­cause it is highly ven­omous, this species is often killed when en­coun­tered by human.

Other Com­ments

The name Both­ri­op­sis bi­lin­eata comes from the Greek words "both­ros" mean­ing "pit" and "-op­sis" mean­ing "face", in ref­er­ence to their fa­cial pit or­gans. The ep­i­thet "bi­lin­eata" is de­rived from Latin. "Bi-" is a pre­fix for "two" and "lin­ea­tus" for "lined or marked with lines". This per­tains to their ven­tro­lat­eral yel­low stripes. Other com­mon names noted for the pitviper are green Tommy goffi, Cobra pa­pa­gaio, lora, orito, and loro machaco. ("Ven­omous An­i­mals and Plants", 2008; Camp­bell and Lamar, 1989; Camp­bell and Lamar, 2004; Tra­vers, 1999)

Con­trib­u­tors

Evin Guil­liams (au­thor), Rad­ford Uni­ver­sity, Brian In­gram (au­thor), Rad­ford Uni­ver­sity, Chris­tine Small (ed­i­tor), Rad­ford Uni­ver­sity, John Berini (ed­i­tor), An­i­mal Di­ver­sity Web Staff.

Glossary

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

arboreal

Referring to an animal that lives in trees; tree-climbing.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

infrared/heat

(as keyword in perception channel section) This animal has a special ability to detect heat from other organisms in its environment.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

ovoviviparous

reproduction in which eggs develop within the maternal body without additional nourishment from the parent and hatch within the parent or immediately after laying.

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

tactile

uses touch to communicate

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

venomous

an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).

vibrations

movements of a hard surface that are produced by animals as signals to others

visual

uses sight to communicate

Ref­er­ences

Rhett A But­ler. 2010. "De­for­esta­tion in the Ama­zon" (On-line). Mongabay.​com. Ac­cessed March 20, 2011 at http://​www.​mongabay.​com/​brazil.​html.

Armed Forces Pest Man­age­ment Board. Ven­omous An­i­mals and Plants. DoD Di­rec­tive 4715.1. Wash­ing­ton D.C.: Armed Forces Pest Man­age­ment Board. 2008. Ac­cessed Feb­ru­ary 03, 2011 at http://​www.​afpmb.​org/​pubs/​living_​hazards/​snakes.​html#​Bothriopsisbilineata.

Al­me­dia-San­tos, S., M. Sa­lo­mao. 2002. Re­pro­duc­tion in neotrop­i­cal pitvipers, with em­pha­sis on species of the genus Both­rops. Bi­ol­ogy of the Vipers, 1: 445-462.

Bar­rio-Amorós, César Luis, 2010. A new Ceuthoman­tis (Anura: Ter­rarana: Ceuthoman­ti­dae) from Saris­ariñama Tepui, South­ern Venezuela. Her­peto­log­ica, 66(2): 172-181.

Camp­bell, J., W. Lamar. 2004. The Ven­omous Rep­tiles of the West­ern Hemi­sphere. United Kin­dom: Comb­stock Pub­lish­ing.

Camp­bell, J., W. Lamar. 1989. Ven­omous Rep­tiles of Latin Amer­ica. Ithaca: Com­stock Pub­lish­ing.

Daly, D., J. Mith­cell. 2000. Low­land Veg­e­ta­tion of Trop­i­cal South Amer­ica -- An Overview. Pp. 391-454 in D Lentz, ed. Im­per­fect Bal­ance: Land­scape Trans­for­ma­tions in the pre-Columbian Amer­i­cas. New York: Co­lum­bia Uni­ver­sity Press.

Fen­wick, Allyson M., G., P. Evans, Jen­nafer A.. 2009. Mor­pho­log­i­cal and mol­e­c­u­lar ev­i­dence for phy­logeny and clas­si­fi­ca­tion of South Amer­i­can pitvipers, gen­era Both­rops, Both­ri­op­sis, and Both­ro­cophias (Ser­pentes: Viperi­dae). Zo­o­log­i­cal Jour­nal of the Lin­nean So­ci­ety, 156/3: 617-640.

Mon­teiro, C., C. Mont­gomery, F. Spina, R. Sawaya, M. Mar­tins. 2006. Feed­ing, re­pro­duc­tion, and mor­phol­ogy of Both­rops mat­togrossen­sis (Ser­pentes, Viperi­dae, Cro­tali­nae) in the Brazil­ian Pan­tanal. Jour­nal of Her­petol­ogy, 40/3: 408–413,.

O'Shea, M. 2005. Ven­omous Snakes of the World. Lon­don: New Hol­land Pub­lish­ers (UK) Ltd.

Parkin­son, Christo­pher L., 1999. Mol­e­c­u­lar sys­tem­at­ics and bio­geo­graph­i­cal his­tory of pitvipers as de­ter­mined by mi­to­chon­dr­ial ri­bo­so­mal DNA se­quences. Copeia, 1999/3: 576-586.

Porto, B., C. Telli, T. Dutra, L. Alves, M. Bozza, C. Fin, F. Thiesen, M. Ren­ner. 2007. Bio­chem­i­cal and bi­o­log­i­cal char­ac­ter­i­za­tion of the ven­oms of Both­ri­op­sis bi­lin­eata and Both­ri­op­sis tae­ni­ata (Ser­pentes: Viperi­dae). Tox­i­con, 50/2: 270-277.

Ro­drigo C. G. de Souza, M.D., , Fun­daçã o Hos­pi­ta­lar Itacare. 2007. A rare ac­ci­dent. Bul­letin of the Chicago Her­peto­log­i­cal So­ci­ety, 42/10: 161-163.

Small­i­gan, R., J. Cole, N. Brito, G. Laing, B. Mertz, S. Manock, J. Maudlin, B. Quist, G. Hol­land, S. Nel­son, D. Lal­loo, G. Ri­vadeneira, M. Bar­ra­gan, D. Dol­ley, M. Ed­dle­ston, D. War­rell, D. Theak­ston. 2004. Cro­taline snake bite in the Ecuado­rian Ama­zon: ran­domised dou­ble blind com­par­a­tive trial of three South Amer­i­can poly­spe­cific an­tiven­oms. British Med­ical Jour­nal, 329: 1129-1133.

Tra­vers, P. 1999. "The Life Cycle of a Pit Viper" (On-line). eHow. Ac­cessed March 19, 2011 at http://​www.​ehow.​com/​about_​6651496_​life-cycle-pit-viper.​html.

Val­dujo, P., N. Cris­tiano, M. Mar­cio. 2002. Ecol­ogy of Both­rops neuwiedi pauloen­sis (Ser­pentes: Viperi­dae: Cro­tali­nae) in the Brazil­ian Cer­rado. Jour­nal of Her­petol­ogy, 36/2: 169-176.

Vidal, N., G. Lecoin­tre, J. Vie, J. Gasc. 1997. Mol­e­c­u­lar sys­tem­at­ics of pitvipers: pa­ra­phyly of the Both­rops com­plex. Life Sci­ences, 320: 95-101.

Vitt, L., J. Cald­well. 1993. Her­petol­ogy. Lon­don, UK: Aca­d­e­mic Press.

Waldez, Fabi­ano, , Vogt, Richard C.. 2009. Eco­log­i­cal and epi­demi­o­log­i­cal as­pects of snakebites in river­side com­mu­ni­ties of the lower Purus River, Ama­zonas, Brazil. Acta Ama­zon­ica, 39/3: 681-692.

Wuster, Wolf­gang, W., Golay, Philippe. 1997. Syn­op­sis of re­cent de­vel­op­ments in ven­omous snake sys­tem­at­ics. Tox­i­con, 35/3: 319-340.