Pit viper

The Crotalinae, commonly known as pit vipers,[2][3] crotaline snakes (named for the Ancient Greek: κρόταλον krotalon[4] castanet/rattle of a rattlesnake's tail), or pit adders, are a subfamily of venomous vipers found in Eurasia and the Americas. They are distinguished by the presence of a heat-sensing pit organ located between the eye and the nostril on both sides of the head. Currently, 22 genera and 151 species are recognized:[5] These are also the only viperids found in the Americas. The groups of snakes represented here include rattlesnakes, lanceheads, and Asian pit vipers. The type genus for this subfamily is Crotalus, of which the type species is the timber rattlesnake, C. horridus.

Pit viper
Timber rattlesnake, Crotalus horridus
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Squamata
Suborder:	Serpentes
Family:	Viperidae
Subfamily:	Crotalinae Oppel, 1811
Synonyms
Crotalini Oppel, 1811 Crotales Cuvier, 1817 Crotalidae Gay, 1825 Crotaloidae Fitzinger, 1826 Cophiadae Boie, 1827 Crotaloidei Eichwald, 1831 Crotalina Bonaparte, 1831 Bothrophes Fitzinger, 1843 Crotalinae Cope, 1860 Teleuraspides Cope, 1871 Crotalida Strauch, 1873 Bothrophera Garman, 1884 Cophiinae Cope, 1895 Lachesinae Cope, 1900 Lachesinii Smith, Smith & Sawin, 1977 Agkistrodontinii Hoge & Romano-Hoge, 1981 Agkistrodontini Hoge & Romano-Hoge, 1983[1]

These snakes range in size from the diminutive hump-nosed viper, Hypnale hypnale, that grows to an average total length (including tail) of only 30–45 cm (12–18 in), to the bushmaster, Lachesis muta, a species known to reach a maximum total length of 3.65 m (12.0 ft) in length.

What makes this subfamily unique is that all member species share a common characteristic: a deep pit, or fossa, in the loreal area between the eye and the nostril on either side of the head. These loreal pits are the external openings to a pair of extremely sensitive infrared-detecting organs, which in effect give the snakes a sixth sense to help them find and perhaps even judge the size of the small, warm-blooded prey on which they feed.[6] Osine triphosphate, monoamine oxidase, generalized esterases and acetylcholine esterase have also been found in it.[6] When prey comes into range, infrared radiation falling onto the membrane allows the snake to determine its direction.[2] Experiments have shown, when deprived of their senses of sight and smell, these snakes can strike accurately at moving objects less than 0.2 °C (0.36 °F) warmer than the background.[7] The paired pit organs provide the snake with thermal rangefinder capabilities.[8] These organs are of great value to a predator that hunts at night, as well as for avoiding the snake's own predators.[9]

Among vipers, these snakes are also unique in that they have a specialized muscle, called the muscularis pterigoidius glandulae, between the venom gland and the head of the ectopterygoid. Contraction of this muscle, together with that of the m. compressor glandulae, forces venom out of the gland.[6]

Geographic range

The subfamily Crotalinae is found in the Old World from eastern Europe eastward through Asia to Japan, China, Indonesia, peninsular India, Nepal, and Sri Lanka. In the Americas, they range from southern Canada southward to Central America to southern South America.[1]

Habitat

The loreal pit is clearly visible below and anterior to the eye (below and posterior to the nostril).

Crotalines are a versatile subfamily, with members found in habitats ranging from parched desert (e.g., the sidewinder, Crotalus cerastes) to rainforests (e.g., the bushmaster, Lachesis muta). They may be either arboreal or terrestrial, and one species is even semiaquatic: the cottonmouth, Agkistrodon piscivorus. The altitude record is held jointly by Crotalus triseriatus in Mexico and Gloydius strauchi in China, both of which have been found above the treeline at over 4,000 m elevation.[6]

Behavior

Although a few species of crotalines are highly active by day, such as Trimeresurus trigonocephalus, a bright green pit viper endemic to Sri Lanka, most are nocturnal, preferring to avoid high daytime temperatures and to hunt when their favored prey are also active. The snakes' heat-sensitive pits are also thought to aid in locating cooler areas in which to rest.

As ambush predators, crotalines typically wait patiently somewhere for unsuspecting prey to wander by. At least one species, the arboreal Gloydius shedaoensis of China, is known to select a specific ambush site and return to it every year in time for the spring migration of birds. Studies have indicated these snakes learn to improve their strike accuracy over time.[10]

Many temperate species of pit vipers (e.g. most rattlesnakes) congregate in sheltered areas or "dens" to overwinter (see hibernation), the snakes benefiting from the combined heat. In cool temperatures and while pregnant, pit vipers also bask on sunny ledges. Some species do not mass together in this way, for example the copperhead, Agkistrodon contortrix, or the Mojave rattlesnake, Crotalus scutulatus.

Like most snakes, crotalines keep to themselves and strike only if cornered or threatened. Smaller snakes are less likely to stand their ground than larger specimens. Pollution and the destruction of rainforests have caused many pit viper populations to decline. Humans also threaten pit vipers, as many are hunted for their skins or killed by cars when they wander onto roads.

Reproduction

With few exceptions, crotalines are ovoviviparous, meaning that the embryos develop within eggs that remain inside the mother's body until the offspring are ready to hatch, at which time the hatchlings emerge as functionally free-living young. In such species the eggshells are reduced to soft membranes that the young shed, either within the reproductive tract, or immediately after emerging.

Among the oviparous (egg-laying) pit vipers are Lachesis, Calloselasma, and some Trimeresurus species. All egg-laying crotalines are believed to guard their eggs.

Brood sizes range from two for very small species, to as many as 86 for the fer-de-lance, Bothrops atrox, which is among the most prolific of all live-bearing snakes.

Many young crotalines have brightly coloured tails that contrast dramatically with the rest of their bodies. Used in a behavior known as caudal luring, the young snakes make worm-like movements with their tails to lure unsuspecting prey within striking distance.

Taxonomy

In the past, the pit vipers were usually classed as a separate family: the Crotalidae. Today, however, the monophyly of the viperines and the crotalines as a whole is undisputed, which is why they are treated here as a subfamily of the Viperidae.

Genera

Genus[ref 1]	Taxon author[ref 1]	Species[ref 1]	Common name	Geographic range[ref 2]
Agkistrodon	Palisot de Beauvois, 1799	6	Moccasins	North America from the northeastern and central USA southward through peninsular Florida and southwestern Texas. In Central America on the Atlantic versant from Tamaulipas and Nuevo León southward to the Yucatán Peninsula, Belize and Guatemala. Along the Pacific coastal plain and lower foothills from Sonora south through Guatemala, El Salvador, Honduras and Nicaragua to northwestern Costa Rica.
Atropoides	Werman, 1992	1	Picado's jumping pitviper	Costa Rica and western Panama.
Bothriechis	Peters, 1859	11	Palm-pitvipers	Southern Mexico (southeastern Oaxaca and the northern highlands of Chiapas), through Central America to northern South America (Colombia, western Venezuela, Ecuador and northern Peru.
Bothrocophias	Gutberlet & Campbell, 2001	6		northern South America.
Bothrops	Wagler, 1824	45	Lanceheads	Northeastern Mexico (Tamaulipas) southward through Central and South America to Argentina; Saint Lucia and Martinique in the Lesser Antilles; Ilha da Queimada Grande off the coast of Brazil.
Calloselasma	Cope, 1860	1	Malayan pitviper	Southeast Asia from Thailand to northern Malaysia and Java, Indonesia.
Cerrophidion	Campbell & Lamar, 1992	5	Montane pitvipers	Southern Mexico (highlands of Guerrero and southeastern Oaxaca), southward through the highlands of Central America (Guatemala, El Salvador, Honduras, northern Nicaragua, Costa Rica) to western Panama.
Crotalus^T	Linnaeus, 1758	51	Rattlesnakes	The Americas, from southern Canada to northern Argentina.
Deinagkistrodon	Gloyd, 1979	1	Hundred-pace pitviper	Southeast Asia.
Garthius	Malhotra & Thorpe, 2004	1	Mount Kinabalu pit viper, Chasen's mountain pit viper	Borneo.
Gloydius	Hoge & Romano-Hoge, 1981	22	Asian moccasins	Russia, east of the Ural Mountains through Siberia, Iran, the Himalayas from Pakistan, India, Nepal and China, Korea, Japan and the Ryukyu Islands.
Hypnale	Fitzinger, 1843	3	Hump-nosed pit vipers	Sri Lanka and India.
Lachesis	Daudin, 1803	4	Bushmasters	Central and South America.
Metlapilcoatlus	Campbell, Frost, & Castoe, 2019	5	jumping pitvipers	The mountains of eastern Mexico southeastward on the Atlantic versant and lowlands though Central America to central Panama. On the Pacific versant, they occur in isolated populations in east-central and southern Mexico, Guatemala, El Salvador, Costa Rica, and Panama.
Mixcoatlus	Jadin, H.M.Smith & Campbell, 2011	3	Mexican pit vipers	Mexico.
Ophryacus	Cope, 1887	3	Mexican horned pitvipers	Mexico.
Ovophis	Burger, 1981	6	Mountain pit vipers	Nepal and Seven Sisters (Assam) of India eastward through Myanmar, Cambodia, Thailand, Laos, Vietnam, West Malaysia, Taiwan, Japan (Okinawa) and Indonesia (Sumatra).
Porthidium	Cope, 1871	9	Hognose pit vipers	Mexico (Colima, Oaxaca and Chiapas on the Pacific side, the Yucatán Peninsula on the Atlantic side) southward through Central America to northern South America (Ecuador in the Pacific lowlands, northern Venezuela in the Atlantic lowlands).
Protobothrops	Hoge & Romano-Hoge, 1983	14	pit vipers	Asia.
Sistrurus	Garman, 1883	2	Ground rattlesnakes	Southeastern Canada, eastern and northwestern USA, isolated populations in northern and central Mexico.
Trimeresurus	Lacépède, 1804	48	Asian lanceheads	Southeast Asia from India to southern China and Japan, and the Malay Archipelago to Timor.
Tropidolaemus	Wagler, 1830	5	Temple vipers	Southern India and Southeast Asia.

*) Not including the nominate subspecies.
^T) Type genus.[ref 2]

1. "Crotalinae". Integrated Taxonomic Information System. Retrieved 26 October 2006.
2. McDiarmid RW, Campbell JA, Touré T. 1999. Snake Species of the World: A Taxonomic and Geographic Reference, vol. 1. Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).

See also

• List of crotaline species and subspecies

References

1. McDiarmid RW, Campbell JA, Touré T (1999). Snake Species of the World: A Taxonomic and Geographic Reference, Volume 1. Washington, District of Columbia: Herpetologists' League. 511 pp. ISBN 1-893777-00-6 (series). ISBN 1-893777-01-4 (volume).
2. Mehrtens JM (1987). Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
3. Sometimes spelled "pitvipers" -- Campbell & Lamar, 2004
4. κρόταλον. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project.
5. "Crotalinae". Integrated Taxonomic Information System. Retrieved 26 October 2006.
6. Campbell JA, Lamar WW (2004). The Venomous Reptiles of the Western Hemisphere. Ithaca and London: Comstock Publishing Associates. 870 pp., 1,500 plates. ISBN 0-8014-4141-2.
7. Parker HW, Grandison AGC (1977). Snakes – A Natural History. Second Edition. London and Ithaca: British Museum (Natural History) and Cornell University Press. 108 pp. 16 plates. LCCCN 76-54625. ISBN 0-8014-1095-9 (cloth), ISBN 0-8014-9164-9 (paper).
8. Bullock TH, Diecke FPJ (1956). "Properties of an infrared receptor". Journal of Physiology 134: 47-87.
9. Stidworthy J (1974). Snakes of the World. New York: Grosset & Dunlap Inc. 160 pp. ISBN 0-448-11856-4.
10. Shine R, Sun L, Kearney M, Fitzgerald M (2002). "Why do Juvenile Chinese Pit-Vipers (Gloydius shedoaensis) Select Arboreal Ambush Sites?" Ethology 108: 897-910. ISSN 0179-1613. PDF Archived 24 July 2008 at the Wayback Machine at University of Sydney School of Biological Sciences. Accessed 26 October 2006.

Further reading

• Gumprecht, Andreas; Tillack, Frank (2004). "A proposal for a replacement name of the snake genus Ermia Zhang, 1993". Russian Journal of Herpetology 11: 73-76.
• Wright, Albert Hazen; Wright, Anna Allen (1957). Handbook of Snakes of the United States and Canada. Ithaca and London: Comstock Publishing Associates, a Division of Cornell University Press. 1,105 pp. (in two volumes). (Seventh Printing 1985). ("Crotalidae", p. 901).
• Goris RC (2011). "Infrared organs of snakes: an integral part of vision". Journal of Herpetology 45: 2-14.

External links

• Pit organs at Life is Short, but Snakes are Long