Bothrops Asper
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The terciopelo (''Bothrops asper'') is a species of
pit viper The Crotalinae, commonly known as pit vipers,Mehrtens JM (1987). ''Living Snakes of the World in Color''. New York: Sterling Publishers. 480 pp. . or pit adders, are a subfamily (biology), subfamily of Viperidae, vipers found in Asia and the ...
s, found from north-east
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
to northern
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. At low to moderate elevations, its habitat ranges from northeastern
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
to
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
n and
Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...
ian
Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...
, as well as
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
. With a mass of up to , and a maximal length of , the terciopelo is among the largest
vipers Vipers are snakes in the family Viperidae, found in most parts of the world, except for Antarctica, Australia, Hawaii, Madagascar, New Zealand, Ireland, and various other isolated islands. They are venomous and have long (relative to non-viper ...
. It is light to dark brown in color, often with yellowish, zig-zag patterning on either side of its body. Dubbed "the ultimate pit-viper" for its defensiveness, large size, fangs and potent venom yield, it has a fearsome reputation, responsible for the most envenomated
snakebites A snakebite is an injury caused by the bite of a snake, especially a venomous snake. A common sign of a bite from a venomous snake is the presence of two puncture wounds from the animal's fangs. Sometimes venom injection from the bite may oc ...
within its range, largely due to its proximity to humans, livestock and pets. Nonetheless, like all
venomous snakes ''Venomous snakes'' are species of the suborder Serpentes that are capable of producing venom, which they use for killing prey, for defense, and to assist with digestion of their prey. The venom is typically delivered by injection using hollow ...
, the terciopelo actively avoids contact with humans and larger animals, with bites generally only occurring when the snake is cornered, pursued, or threatened. No subspecies are currently recognized.


Etymology

The
generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...
name, ''Bothrops'', comes from the Greek words and , which mean 'pit' and 'face' (or 'eye'), respectively. This is a reference to these snakes' highly sensitive heat-detecting pit organs. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
, , which is a Latin word meaning 'rough' or 'harsh', may allude to the species' keeled dorsal scales.


Common names

Some of the common names applied to this snake are ''terciopelo'' ("velvet" in Spanish), ''fer-de-lance'', ''mapepire balsain'' (Trinidad), ''carpet labaria'' (Guyana), ''barba amarilla'' (Guatemala, Honduras, Nicaragua; "yellow beard"), ''equis'' (Ecuador and Panama; "x"), ''talla equis'' , ''mapaná'' (Colombia), mapanare (Venezuela), ''nauyaca'' (México; from
Nahuatl Nahuatl ( ; ), Aztec, or Mexicano is a language or, by some definitions, a group of languages of the Uto-Aztecan language family. Varieties of Nahuatl are spoken by about Nahuas, most of whom live mainly in Central Mexico and have smaller popul ...
''nahui'', four, and ''yacatl'', nose; "four noses"), and ''yellow-jaw tommygoff'' (Belize)." The name fer-de-lance is French (or possibly Créole), translating to "iron of the lance", "iron spear point" or simply "spearhead" or "lancehead".Lillywhite, Harvey B. (2008). Dictionary of Herpetology. Krieger Publishing Co. Malabar, Florida. viii, 376 pp. Greene, Harry W. (1997). Snakes: The Evolution of Mystery in Nature. University of California Press, Berkeley, California. xiii, 351 pp. In English the name lancehead is applied to the genus ''
Bothrops ''Bothrops'' is a genus of highly venomous pit vipers Endemism, endemic to the Neotropics.McDiarmid RW, Jonathan A. Campbell, Campbell JA, Touré T. 1999. ''Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1''. Herpetologist ...
'' as a whole,Tipton, Bob L. (2005). Snakes of the Americas, Checklist and Lexicon. Krieger Publishing Co. Malabar, Florida. xiv, 477 pp. (With CD) and in combination with the majority of the species within the genus (e.g.: Alcatrazes lancehead (''
Bothrops alcatraz ''Bothrops alcatraz'', also known as the Alcatrazes lancehead, is a pitviper species found only on the Alcatrazes Islands off the coast of southeastern Brazil. No subspecies are currently recognized. Like all pit vipers, it is venomous. Descr ...
''), Patagonian lancehead, (''
Bothrops ammodytoides :''Common names: Patagonian lancehead,Campbell JA, Lamar WW. 2004. ''The Venomous Reptiles of the Western Hemisphere''. Comstock Publishing Associates, Ithaca and London. 870 pp. 1500 plates. . yarará ñata,Bothrops andianus'') etc.). The name fer-de-lance has been used inconsistently and very informally with several species in the genus ''Bothrops'' having been called fer-de-lance at one time or another, most commonly ''Bothrops asper'', ''
Bothrops atrox ''Bothrops atrox'' — also known as the common lancehead, ''fer-de-lance'', ''barba amarilla'',Campbell JA, Lamar WW. 2004. ''The Venomous Reptiles of the Western Hemisphere''. Comstock Publishing Associates, Ithaca and London. 870 pp. 1500 plat ...
'', ''
Bothrops caribbaeus ''Bothrops'' is a genus of highly venomous pit vipers endemic to the Neotropics.McDiarmid RW, Campbell JA, Touré T. 1999. ''Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1''. Herpetologists' League. 511 pp. (series). ...
'', and ''
Bothrops lanceolatus ''Bothrops lanceolatus'' — known as the ''fer-de-lance'', Martinican pit viper,Brown JH. 1973. ''Toxicology and Pharmacology of Venoms from Poisonous Snakes''. Springfield, Illinois: Charles C. Thomas. 184 pp. LCCCN 73-229. . and Martinique lan ...
''.Schwartz, Albert and Robert W. Henderson. (1991). ''Amphibians and Reptile of the West Indies: Descriptions, Distributions, and Natural History''. University of Florida Press. Gainesville, Florida. xvi, 720 pp. Henderson, Robert W. and Robert Powell. (2009). ''Natural History of West Indian Reptiles and Amphibians''. University Press of Florida. Gainesville, Florida. xxiv, 495 pp. Several herpetologists have preferred and argued to restrict the name fer-de-lance to the Martinique lancehead (''
Bothrops lanceolatus ''Bothrops lanceolatus'' — known as the ''fer-de-lance'', Martinican pit viper,Brown JH. 1973. ''Toxicology and Pharmacology of Venoms from Poisonous Snakes''. Springfield, Illinois: Charles C. Thomas. 184 pp. LCCCN 73-229. . and Martinique lan ...
'') from the French speaking
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
Island of
Martinique Martinique ( ; or ; Kalinago language, Kalinago: or ) is an island in the Lesser Antilles of the West Indies, in the eastern Caribbean Sea. It was previously known as Iguanacaera which translates to iguana island in Carib language, Kariʼn ...
, but popular usage has rarely recognized any distinction. In their seminal opus on the venomous reptiles of the Western hemisphere, Campbell & Lamar stated: "The name fer-de lance, widely used in North America with reference to ''B. asper'' and ''B. atrox'', has no legitimate origin of use in regions inhabited by this snake" .g. Spanish and Portuguese speaking countries In an effort to establish standardized names for the amphibians and reptiles of North America, the
Society for the Study of Amphibians and Reptiles The Society for the Study of Amphibians and Reptiles (SSAR) is an international herpetological society. It is a non-profit organization supporting education, conservation, and research related to reptiles and amphibians. Regular publications ...
(SSAR), a not-for-profit organization and one of the largest international herpetological societies, applied the following nomenclature: * ''Bothrops'' Wagler, 1824: Lanceheads (English) * ''Bothrops asper'' (Garman, 1884
883 __NOTOC__ Year 883 ( DCCCLXXXIII) was a common year starting on Tuesday of the Julian calendar. Events By place Europe * Spring – Viking raiders ravage Flanders, and sack the abbey at Saint-Quentin. King Carloman II blocks thei ...
: Terciopelo (English) In recent decades, herpetologists have preferred the name 'terciopelo' for ''Bothrops asper'',Lee, Julian C. (2000). A Field Guide to the Amphibians and Reptiles of the Maya World. Comstock Publishing Associates, Cornell University Press, Ithaca, New York. xi + 402 pp. Savage, Jay M. (2002). The Amphibians and Reptiles of Costa Rica, A Herpetofauna between Two Continents, between Two Seas. University of Chicago Press, Chicago, Illinois. xx, 934 pp. Köhler, Gunther (2008). Reptiles of Central America, 2nd Edition. Herpeton, Verlag Elke Köhler, Offenbach, Germany. 400 pp. Lemos Espinal, Julio A. and James R. Dixon. (2013). Amphibians and Reptiles of San Luis Potosí. Eagle Mountain Publishing, LC. Eagle Mountain, Utah. i-xii, 1-300 pp. although the term fer-de-lance is still common in popular usage.


Description

''Bothrops'' species can be distinguished by their broad, flattened heads which are set apart from the rest of their bodies. The head of this snake is light to dark brown or even black. Although usually absent, it may have occipital blotches or streaks that range from indistinct to distinct. The underside is most often pale yellow. This species has different patterns and colors on its dorsal and ventral sides and it exhibits a postorbital stripe. The ventral side is yellow, cream, or a whitish gray, with dark blotches that are more frequent closer to the posterior end. Ventrolaterally, ''B. asper'' has interchanging gray scales which are more pale towards the medial line. Dark triangles with pale edges can be seen laterally, which range in number from 18 to 25. Apices either alternate or are reflective of each other over the middorsal line. In the interspaces, there are dark, paravertebral blotches. Specimens may have a yellow zig-zag-shaped line on each side of the body. There is a great variety of colours on its dorsal side: olive, gray, light brown to dark brown, tan or sometimes nearly black. To prevent water loss where they occur in drier regions, this species has more scales. Specimens of this species may weigh up to and are often in length. Very big females can reach lengths up to , although this is uncommon. These are among the most
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
of all snakes. The two sexes are born the same size, but by age 7 to 12 months, females begin to grow at a much faster rate than males. Females have thick, heavy bodies and grow significantly larger than males. They also have heads two or three times the size of males relative to their size and proportionally bigger fangs (typically 2.5 cm), as well. Across its geographic range, this species varies greatly phenotypically. As a result, great confusion between it and other related species, most notably ''Bothrops atrox'', which is similar in color but usually has yellow or rust-like tones and rectangular or trapezoidal blotches.


Distribution

It is found on the Gulf - Atlantic versant of eastern Mexico as far north as the state of Tamaulipas, southward through the entire Yucatán Peninsula extending into Central America, including Guatemala, Belize, Honduras, Nicaragua, Costa Rica and Panama. An isolated population occurs in southeastern Chiapas (Mexico) and southwestern Guatemala. In northern South America, it is found in Colombia, Ecuador, Guyana and Venezuela. The type locality given is "Obispo, on the
Isthmus of Darien The Isthmus of Panama, historically known as the Isthmus of Darien, is the narrow strip of land that lies between the Caribbean Sea and the Pacific Ocean, linking North and South America. The country of Panama is located on the isthmus, along w ...
" (Panama).Farr, William L. and David Lazcano. 2017. Distribution of Bothrops asper in Tamaulipas, Mexico and a review or prey items. Southwestern Naturalist 62(1): 77-84. This is mostly a lowland species that, in Mexico and Central America, occurs from about sea level to altitude. In South America, it apparently ranges to considerably higher elevations: up to in Venezuela and at least in Colombia according to herpetologist Lancini. According to Campbell and Lamar (2004), its range in Ecuador extends as far south along the Pacific coast as
El Oro Province El Oro (; ''oro'' = gold) is the southernmost of Ecuador's coastal provinces. It was named for its historically important gold production. Today it is one of the world's major exporters of bananas. The capital is Machala. History The area was s ...
and the Vilcabamba area of the Río Catamayo Valley. This species is reported to occur from seven (Bolívar, Carchi, Chimborazo, Esmeraldas, Guayas, Los Ríos and Pichincha) of the fourteen provinces along the Pacific slope of Ecuador. There are even a few records from northern coastal
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
, with these snakes being reported in the
Tumbes Region Tumbes () is a coastal department and region in northwestern Peru bordering Ecuador. Due to the region's location near the Equator it has a warm climate, with beaches that are considered among the finest in Peru. Despite its small area, the re ...
. It is also known from the island of Gorgona off the Pacific coast of
Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...
. ''B. asper'' occurs throughout the inter-
Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...
valleys of Colombia across the Caribbean coastal plain through central Venezuela north of the
Orinoco The Orinoco () is one of the longest rivers in South America at . Its drainage basin, sometimes known as the Orinoquia, covers approximately 1 million km2, with 65% of it in Venezuela and 35% in Colombia. It is the List of rivers by discharge, f ...
as far east as the
Delta Amacuro Delta Amacuro State (, ) is one of the 23 States of Venezuela, states of Venezuela, and is the location of the Orinoco Delta. The Paria Gulf and the Atlantic Ocean are found to the north, Bolívar State (Venezuela), Bolívar State is found to th ...
region. This is the only ''
Bothrops ''Bothrops'' is a genus of highly venomous pit vipers Endemism, endemic to the Neotropics.McDiarmid RW, Jonathan A. Campbell, Campbell JA, Touré T. 1999. ''Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1''. Herpetologist ...
'' species that occurs on the island of
Trinidad Trinidad is the larger, more populous island of the Republic of Trinidad and Tobago, the country. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is the southernmost island in ...
, although the situation there is complicated due to proximity of Trinidad to the
Orinoco Delta The Orinoco Delta is a vast river delta of the Orinoco River, located in eastern Venezuela. Location The Orinoco Delta is one of the eight natural regions of Venezuela. It covers the whole of Delta Amacuro State and a few square kilometers o ...
where it may be
sympatric In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. An initially interbreeding population that splits into two or more distinct spe ...
with ''B. atrox''. Due to the casual and informal application of the name "Fer-de-lance" being applied to any number of species of Latin America pit vipers in the genus ''
Bothrops ''Bothrops'' is a genus of highly venomous pit vipers Endemism, endemic to the Neotropics.McDiarmid RW, Jonathan A. Campbell, Campbell JA, Touré T. 1999. ''Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1''. Herpetologist ...
'', there is much confusion and misunderstanding, particularly in popular literature, as to proper nomenclature. Populations of ''Bothrops'' often referred to as Fer-de-lance on the island of
Saint Lucia Saint Lucia is an island country of the West Indies in the eastern Caribbean. Part of the Windward Islands of the Lesser Antilles, it is located north/northeast of the island of Saint Vincent (Saint Vincent and the Grenadines), Saint Vincent ...
are ''
Bothrops caribbaeus ''Bothrops'' is a genus of highly venomous pit vipers endemic to the Neotropics.McDiarmid RW, Campbell JA, Touré T. 1999. ''Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1''. Herpetologists' League. 511 pp. (series). ...
''. Populations of ''Bothrops'' referred to as Fer-de-lance on the island of
Martinique Martinique ( ; or ; Kalinago language, Kalinago: or ) is an island in the Lesser Antilles of the West Indies, in the eastern Caribbean Sea. It was previously known as Iguanacaera which translates to iguana island in Carib language, Kariʼn ...
are regarded as ''
Bothrops lanceolatus ''Bothrops lanceolatus'' — known as the ''fer-de-lance'', Martinican pit viper,Brown JH. 1973. ''Toxicology and Pharmacology of Venoms from Poisonous Snakes''. Springfield, Illinois: Charles C. Thomas. 184 pp. LCCCN 73-229. . and Martinique lan ...
''.


Habitat

This species likes moist environments, and occurs in most life zones located at low or middle elevations (up to ), excluding those with strong seasonal dry periods. They are, however, sometimes found at much higher elevations. This is true in the premontane forest in Costa Rica, the cloud forest of Guatemala and Mexico, or the lower montane wet forest in the Caribbean Region of Colombia and Ecuador. It chiefly inhabits
tropical rainforest Tropical rainforests are dense and warm rainforests with high rainfall typically found between 10° north and south of the Equator. They are a subset of the tropical forest biome that occurs roughly within the 28° latitudes (in the torrid zo ...
and
evergreen forest An evergreen forest is a forest made up of evergreen trees. They occur across a wide range of climatic zones, and include trees such as conifers and holly in cold climates, eucalyptus, live oak, acacias, magnolia, and banksia in more temperate zo ...
, but it also occurs in drier areas of tropical deciduous forest, thorn forest and pine savannah near lakes, rivers and streams. The home range of ''B. asper'' averages between 3.71 ha and 5.95 ha, which is comparatively small in relation to other pitvipers.


Behavior

''B. asper'' is
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
and solitary. It is less active in colder and drier periods. This species is often found near rivers and streams, basking in the sun during the day and lying still while well camouflaged in leaf litter or under forest cover waiting to ambush prey such as rats and mice that come within range during the night. When cornered or threatened, this species can be very defensive and may exhibit an S-coiled defense display. Juveniles are often semiarboreal, and even adults are sometimes encountered in bushes and low trees. Juveniles are also known to exhibit
caudal luring Caudal luring is a form of aggressive mimicry characterized by the waving or wriggling of the predator's tail to attract prey. This movement attracts small animals who mistake the tail for a small worm or other small animal. When the animal approac ...
, a use of their differently colored tail tips to lure prey. Although both males and females display this behavior, only males have bright coloured tail tips. Compared to the common lancehead, ''B. atrox'', these snakes have been described as excitable and unpredictable when disturbed. They can, and often will, move very quickly, usually opting to flee from danger, but are capable of suddenly reversing direction to vigorously defend themselves. Adult specimens, when cornered and fully alert, are dangerous. In a review of bites from this species suffered by field biologists, Hardy (1994) referred to it as the "ultimate pit viper".


Diet

''Bothrops asper'' is a diet generalist and is known to prey on a remarkably wide range of animals. A generalized
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
diet shift occurs, with a higher percentage of
ectothermic An ectotherm (), more commonly referred to as a "cold-blooded animal", is an animal in which internal physiological sources of heat, such as blood, are of relatively small or of quite negligible importance in controlling body temperature.Daven ...
prey in juveniles, changing to a greater percentage of
endotherm An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inst ...
ic prey in adults, particularly small mammals. However both juveniles and adults, regardless of size or age, are known to opportunistically prey on ectothermic and endothermic species. Reports of invertebrate and insect remains in the digestive tracts along with frog and lizard remains are believed to represent secondary ingestion, however the dissection of several specimens containing only insect remains such as beetles (
Coleoptera Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
), and bugs (
Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...
) are believed to reflect insects as primary prey too. Cannibalism has been reported in both captive and wild juveniles and the species is known to scavenge on dead frogs and rodents. Just a few of the documented ectothermic prey items include: small to moderately-sized centipedes (specifically '' Scolopendra angulata''), beetles (
Coleoptera Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
), grasshoppers (
Orthoptera Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – gras ...
), flies (
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advance ...
), hemipterans (
Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...
), ants ( Formicidae), crayfish (
Astacidea Astacidea is an infraorder of decapod crustaceans including lobsters (but not "lobsters" such as the spiny lobster etc.), crayfish, and their close relatives. Description The Astacidea are distinguished from most other decapods by the presence ...
), eels (''
Synbranchus ''Synbranchus'' is a genus of swamp eels native to Central and South America. Species There are currently four recognized species in this genus: * '' Synbranchus lampreia'' Favorito, Zanata & Assumpção, 2005 * '' Synbranchus madeirae'' D. ...
''); caecilians (''
Dermophis ''Dermophis'' is a genus of worm-like amphibians in the family Dermophiidae, the Neotropical and Tropical African caecilians. They are found in the Middle America between southern Mexico and northwestern Colombia Colombia, officially th ...
''), frogs (''
Eleutherodactylus ''Eleutherodactylus'' is a genus of frogs in the family Eleutherodactylidae.Hedges, S. B., W. E. Duellman, and M. P. Heinicke . 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and c ...
'', ''
Leptodactylus ''Leptodactylus'' is a genus of leptodactylid frogs. It includes the species commonly called ditch frogs or white-lipped frogs. It is very similar to ''Physalaemus'', a close relative, and indeed the 2005 described ''Leptodactylus lauramiriamae' ...
'', ''
Lithobates ''Lithobates'', commonly known as the bullfrogs, is a genus of true frogs, of the family Ranidae. The name is derived from '' litho-'' (stone) and the Greek ' (, one that treads), meaning one that treads on rock, or rock climber. As presently de ...
'', ''
Pristimantis ''Pristimantis'' is a very specious genus of frogs distributed in the southern Caribbean islands (Lesser Antilles) and in Central and South America from Honduras to northern Argentina and southern Brazil. With 617 described species (as of Februa ...
'', ''
Rhinella ''Rhinella'', commonly known as South American toads, beaked toads or Rio Viejo toads, is a genus of true toads native to Neotropical parts of Mexico, Central and South America. Additionally, the cane toad has been Introduced species, Introduced ...
'', '' Smilisca''), toads (''
Rhinella ''Rhinella'', commonly known as South American toads, beaked toads or Rio Viejo toads, is a genus of true toads native to Neotropical parts of Mexico, Central and South America. Additionally, the cane toad has been Introduced species, Introduced ...
''), amphisbaenians (''
Amphisbaena The amphisbaena (, , or , plural: amphisbaenae; ) is a mythological, ant-eating serpent with a head at each end. The name of the creature is alternatively written amphisbaina, amphisbene, amphisboena, amphisbona, amphista, amfivena, amphivena, ...
''), lizards (''
Alopoglossus ''Alopoglossus'' is a genus of lizards in the monogeneric Family (biology), family Alopoglossidae. The genus is distributed from Costa Rica in Central America to northern South America. Species The genus ''Alopoglossus'' contains the following 3 ...
'', ''
Ameiva ''Ameiva'', commonly called jungle-runners, is a genus of whiptail lizards that belongs to the family Teiidae. Geographic range Member species of the genus ''Ameiva'' are found in South America, Central America and the Caribbean (West Indies). ...
'', ''
Anolis ''Anolis'' is a genus of anoles (), iguanian lizards in the family Dactyloidae, native to the Americas. With more than 425 species, it represents the world's most species-rich amniote tetrapod genus, although many of these have been proposed to b ...
'', ''
Ctenosaura ''Ctenosaura'' is a lizard genus commonly known as spinytail iguanas or ctenosaurs. The genus is part of the large lizard family Iguanidae and is native to Mexico and Central America. The name is derived from two Greek words: (), meaning "com ...
'', ''
Gonatodes ''Gonatodes'' is a genus of New World dwarf geckos of the family Sphaerodactylidae. Description The majority of the species in the genus ''Gonatodes'' are diurnally active, scansorial, and sexually dichromatic, with adult body size (snout–ve ...
''), and snakes (''
Bothrops ''Bothrops'' is a genus of highly venomous pit vipers Endemism, endemic to the Neotropics.McDiarmid RW, Jonathan A. Campbell, Campbell JA, Touré T. 1999. ''Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1''. Herpetologist ...
'', ''
Dipsas ''Dipsas'' is a genus of nonvenomous New World snakes in the subfamily Dipsadinae of the family Colubridae. The genus ''Sibynomorphus'' has been moved here. Species of the genus ''Dipsas'' are known as snail-eaters. Taxonomy The genus ''Dipsas'' ...
'', ''
Erythrolamprus ''Erythrolamprus'' is a genus of colubrid snakes native to Central America, the Caribbean, and South America. They include the false coral snakes, which appear to be coral snake mimics. Classification The genus ''Erythrolamprus'' belongs to ...
'', ''
Ninia ''Ninia'', commonly referred to as coffee snakes, is a genus of snakes in the family Colubridae. The genus consists of 12 species that are native to south-eastern Mexico, Central America, and the northern part of South America. Some species are ...
''). Endothermic prey species include: bay wren ('' Cantorchilus nigricapillus''), grey-headed tanager (''
Eucometis penicillata The grey-headed tanager (''Eucometis penicillata'') is a widely distributed species of small Neotropical bird in the tanager family Thraupidae. It is the only member of the genus ''Eucometis''. It is found in Central America and northern South A ...
''), wren ('' Troglodytes''), blue-black grassquit ('' Volatinia jacarina''), Central American woolly opossum (''
Caluromys derbianus Derby's woolly opossum (''Caluromys derbianus''), or the Central American woolly opossum, is an opossum found in deciduous and moist evergreen forests of Central America, from southern Mexico to western Ecuador and Colombia. It was first describ ...
''), common opossum (''
Didelphis marsupialis The common opossum (''Didelphis marsupialis''), also called the southern or black-eared opossum or gambá, and sometimes called a possum, is a marsupial species living from the northeast of Mexico to Bolivia (reaching the coast of the South Paci ...
''), Desmarest's spiny pocket mouse (''
Heteromys desmarestianus ''Heteromys'' is a genus of rodents in the family Heteromyidae, commonly known as spiny pocket mice. It is the only extant genus in the subfamily Heteromyinae which also includes the extinct genera ''Diprionomys'' and ''Metaliomys''. ''Heteromys ...
''), dusky rice rat (''
Melanomys caliginosus ''Melanomys caliginosus'', also known as the dusky melanomysMusser and Carleton, 2005 or dusky rice rat, is a species of rodent in the genus '' Melanomys'' of family Cricetidae. It is found from Central America, in Honduras, Nicaragua, Costa Rica ...
''), black rat (''
Rattus rattus The black rat (''Rattus rattus''), also known as the roof rat, ship rat, or house rat, is a common long-tailed rodent of the stereotypical rat genus ''Rattus'', in the subfamily Murinae. It likely originated in the Indian subcontinent, but is ...
''), Rothschild's porcupine ('' Coendou rothschildi''), Brazilian cottontail (''
Sylvilagus brasiliensis The common tapeti (''Sylvilagus brasiliensis''), also known as the Brazilian cottontail, forest cottontail, or (formerly) simply tapeti is a species of cottontail rabbit. It is small to medium-sized with a small, dark tail, short hind feet, and s ...
''), and least shrew ('' Cryptotis parva'').


Reproduction

The timing of the reproductive cycle and the litter size of this species vary according to location: in some parts of Costa Rica, for example, it is more prolific than in others. Reproduction is highly seasonal and in Costa Rica, reproductive cycles are tightly related to rainfall patterns. The timing of breeding differs between populations in the Caribbean and Pacific lowlands. On the Pacific side, mating took place between September and November, with females giving birth between April and June. The average number of offspring was 18.6 (five to 40) in this population. Neonates ranged in total length from and in weight from . On the Atlantic side, mating was observed in March, and births occurred between September and November. The average number of offspring was 41.1 (14–86), whereas the total length of neonates ranged from , and weighed from . In both populations, gestation time ranged from six to eight months, and the size of a litter correlated significantly with the size of the female. This species is considered to be the most prolific of all snakes in the Americas. Male-male combat in this species has not been observed. Females will mate with more than one male during mating season. Mating includes a series of movements of the male, which then slowly chases an accepting female. The female then stops movement and extends her posture to mate. It is not known whether this species exhibits annual or biannual reproduction.


Venom

''B. asper'', together with ''
Crotalus durissus ''Crotalus durissus'', known as the South American rattlesnake, tropical rattlesnake,Mehrtens JM. 1987. Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. and by #Common names, other names, is a highly venomous snake, ve ...
'', is the leading cause of snakebite in
Yucatán Yucatán, officially the Free and Sovereign State of Yucatán, is one of the 31 states which, along with Mexico City, constitute the 32 federal entities of Mexico. It comprises 106 separate municipalities, and its capital city is Mérida. ...
, Mexico. It is considered the most dangerous snake in Costa Rica, responsible for 46% of all bites and 30% of all hospitalized cases; before 1947, the fatality rate was 9%, but this has since declined to almost 0% (Bolaños, 1984), mostly due to the Clodomiro Picado Research Institute, responsible for the production of snake antiophidic sera (which are also exported to other countries in Latin America and Africa) and scientific research on serpents and their venoms, as well as educational and extension programs in rural areas and hospitals. In the Colombian states of
Antioquia Antioquia is the Spanish form of Antioch. Antioquia may also refer to: * Antioquia Department Antioquia () is one of the 32 departments of Colombia, located in the central northwestern part of Colombia with a narrow section that borders th ...
and Chocó, it causes 50–70% of all snakebites, with a
sequelae A sequela (, ; usually used in the plural, sequelae ) is a pathological condition resulting from a disease, injury, therapy, or other trauma. Derived from the Latin word meaning "sequel", it is used in the medical field to mean a complication or c ...
rate of 9% and a fatality rate of 6% (Otero et al., 1992). In the state of
Lara Lara may refer to: People * Lara (name), can be a given name or a surname in several languages * Lara (mythology), a naiad nymph, daughter of the river Almo in Ovid's ''Fasti'' Places *Lara (state), a state in Venezuela * Electoral district ...
, Venezuela, it is responsible for 78% of all envenomations and all snakebite fatalities (Dao-L., 1971). One of the reasons so many people are bitten is because of its association with human habitation; many bites occur indoors (Sasa & Vázquez, 2003).
Herpetologist Herpetology (from Ancient Greek ἑρπετόν ''herpetón'', meaning "reptile" or "creeping animal") is a branch of zoology concerned with the study of amphibians (including frogs, salamanders, and caecilians (Gymnophiona)) and reptiles (in ...
Douglas March died after being bitten by this species.Warrell DA. 2004. Snakebites in Central and South America: Epidemiology, Clinical Features, and Clinical Management. ''In'' Campbell JA, Lamar WW. 2004. The Venomous Reptiles of the Western Hemisphere. Comstock Publishing Associates, Ithaca and London. 870 pp. 1500 plates. . This species is irritable and fast-moving. It is also regarded as being more excitable and unpredictable than ''B. atrox''. Its large size and habit of raising its head high off the ground can result in bites above the knee. It has also been observed to eject venom over a distance of at least 6 ft (1.8 m) in fine jets from the tips of its fangs (Mole, 1924). Bite symptoms include pain, oozing from the puncture wounds, local swelling that may increase for up to 36 hours, bruising that spreads from the bite site, blisters, numbness, mild fever, headache, bleeding from the nose and gums,
hemoptysis Hemoptysis or haemoptysis is the discharge of blood or blood-stained sputum, mucus through the mouth coming from the bronchi, larynx, vertebrate trachea, trachea, or lungs. It does not necessarily involve coughing. In other words, it is the airw ...
, gastrointestinal bleeding,
hematuria Hematuria or haematuria is defined as the presence of blood or red blood cells in the urine. "Gross hematuria" occurs when urine appears red, brown, or tea-colored due to the presence of blood. Hematuria may also be subtle and only detectable with ...
,
hypotension Hypotension, also known as low blood pressure, is a cardiovascular condition characterized by abnormally reduced blood pressure. Blood pressure is the force of blood pushing against the walls of the arteries as the heart pumps out blood and is ...
, nausea, vomiting, impaired consciousness and tenderness of the
spleen The spleen (, from Ancient Greek '' σπλήν'', splḗn) is an organ (biology), organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The spleen plays important roles in reg ...
. In untreated cases, local necrosis frequently occurs and may cause gangrene which often requires amputation. In 12 fatal cases, the cause of death was
sepsis Sepsis is a potentially life-threatening condition that arises when the body's response to infection causes injury to its own tissues and organs. This initial stage of sepsis is followed by suppression of the immune system. Common signs and s ...
(5),
intracranial hemorrhage Intracranial hemorrhage (ICH) refers to any form of Hemorrhage, bleeding Internal bleeding, within the Human skull, skull. It can result from trauma, vascular abnormalities, hypertension, or other medical conditions. ICH is broadly categorized ...
(3),
acute kidney injury Acute kidney injury (AKI), previously called acute renal failure (ARF), is a sudden decrease in renal function, kidney function that develops within seven days, as shown by an increase in serum creatinine or a decrease in urine output, or both. ...
with
hyperkalemia Hyperkalemia is an elevated level of potassium (K+) in the blood. Normal potassium levels are between 3.5 and 5.0 mmol/L (3.5 and 5.0 mEq/L) with levels above 5.5mmol/L defined as hyperkalemia. Typically hyperkalemia does not cause symptoms. Oc ...
and
metabolic acidosis Metabolic acidosis is a serious electrolyte disorder characterized by an imbalance in the body's acid-base balance. Metabolic acidosis has three main root causes: increased acid production, loss of bicarbonate, and a reduced ability of the kidn ...
(2) and
hemorrhagic shock Hypovolemic shock is a form of shock caused by severe hypovolemia (insufficient blood volume or extracellular fluid in the body). It can be caused by severe dehydration or blood loss. Hypovolemic shock is a medical emergency; if left untreated, ...
(1). Venom yield (dry weight) averages 458 mg, with a maximum of 1530 mg (Bolaños, 1984) and an in mice of 2.844 mg/kg IP. The venomous bite of ''B. asper'' has been suggested to have been a factor in the choice of certain
Mayan Mayan most commonly refers to: * Maya peoples, various indigenous peoples of Mesoamerica and northern Central America * Maya civilization, pre-Columbian culture of Mesoamerica and northern Central America * Mayan languages, language family spoken ...
settlements, such as
Nim Li Punit Nim Li Punit () is a Maya Classic Period site in the Toledo District of the nation of Belize, located 50 kilometres north of the town of Punta Gorda, and directly adjacent to the village of Indian Creek. Nim Li Punit is sometimes known as Big ...
, Belize Central America, where the thick jungle inhabited by these snakes was used as a defensive boundary.Nim Li Punit
by C. Michael Hogan, a
Megalithic Portal
Accessed 15 March 2008.
The venom of the fer-de-lance is so potent that didelphine opossums (i.e., opossums like the
Virginia opossum The Virginia opossum (''Didelphis virginiana''), also known as the North American opossum, is a member of the opossum family found from southern Canada to northern Costa Rica, making it the northernmost marsupial in the world and the only marsup ...
), which are normally immune to the venom of pit vipers and rattlesnakes, are still capable of succumbing to the venom of this snake. This is especially the case if the opossum is not fully grown (and thus the venom is more concentrated per gram). Indeed, the extremely potent venom of ''B. asper'' has been suggested to have evolved as part of an
evolutionary arms race In evolutionary biology, an evolutionary arms race is an ongoing struggle between competing sets of co-evolving genes, phenotypic and behavioral traits that develop escalating adaptations and counter-adaptations against each other, resembling the ...
between these snakes and didelphine opossums, as a defensive adaptation to prevent predation by opossums, an adaptation that allowed fer-de-lances to prey on large opossums, or both.


Taxonomy

This species was once regarded as a subspecies of ''B. atrox'' and can still often be confused with it.


Notes


References


Further reading

* Bolaños R. 1984. ''Serpientes, venenos, y ofidismo en Centroamérica''. Editoria Universidad de Costa Rica, San José. 136 pp. * Dao-L. L. 1971.'' Emponzoñamiento ofícido en el Estado Lara''. Gaceta Medica de Caracas 79: 383–410. * Garman S. 1884 1883" ''The Reptiles and Batrachians of North America''. Memoirs of the Museum of Comparative Zoölogy at Harvard College, Cambridge, Mass. Vol. VIII, No. 3. xxxi + 185 pp. + 10 Plates. (''Trigonocephalus asper'', p. 124.) * Otero R, Tobón GS, Fernando Gómez L, Osorio R, Valderrama R, Hoyos D, Urreta JE, Molina S, Arboleda JJ. 1992. ''Accidente ofídico en Antioquia y Chocó. Aspectos clínicos y epidimiológicos (marzo de 1989-febrero de 1990)''. Acta Médica Colombiana 17: 229–249. * Sasa M, Vázquez S. 2003. Snakebite envenomation in Costa Rica: a revision of incidence in the decade 1990–2000. Toxicon 41(1): 19–22.


External links

* * https://www.iucnredlist.org/species/197464/2486766 {{Taxonbar, from=Q384987 asper Snakes of Central America Reptiles of Mexico Reptiles of Ecuador Reptiles of Guatemala Reptiles of Honduras Reptiles of Panama Reptiles of Colombia Reptiles of Costa Rica Reptiles described in 1884