Crotalinae

Life-forms

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Viperoidea

Familia: Viperidae
Subfamilia: Crotalinae

Genera (21): Agkistrodon – Bothriechis – Bothrocophias – Bothrops – Calloselasma – Cerrophidion – Crotalus – Deinagkistrodon – Garthius – Gloydius – Hypnale – Lachesis – Metlapilcoatlus – Mixcoatlus – Ophryacus – Ovophis – Porthidium – Protobothrops – Sistrurus – Trimeresurus – Tropidolaemus
Name

Crotalinae Oppel, 1811
References
Primary references

Oppel, N. 1811. Die Ordnungen, Familien und Gattungen der Reptilien, etc. Lindauer: Munich.

Additional references

Parkinson, C.L. 1999. Molecular Systematics and Biogeographical History of Pitvipers as Determined by Mitochondrial Ribosomal DNA Sequences. Copeia 1999(3): 576–586. DOI: 10.2307/1447591 Hybrid open access journal Reference page.
Campbell, J.A. & Lamar, W.W. 2004. The Venomous Reptiles of the Western Hemisphere. 2 Volumes. Cornell University Press: Ithaca, New York. 1032 pp. ISBN 9780801441417 Reference page.
Castoe, T.A. & Parkinson, C.L. 2006. Bayesian mixed models and the phylogeny of pitvipers (Viperidae: Serpentes). Molecular Phylogenetics and Evolution 39(1): 91–110. DOI: 10.1016/j.ympev.2005.12.014 Paywall Reference page.
Fenwick, A.M., Gutberlet Jr., R.L., Evans, J.A. & Parkinson, C.L. 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. DOI: 10.1111/j.1096-3642.2008.00495.x.Reference page.
Guo, P. & Wang, Y. 2011. A new genus and species of cryptic Asian green pitviper (Serpentes: Viperidae: Crotalinae) from southwest China. Zootaxa 2918: 1–14. Preview
Alencar, L.R.V., Quental, T.B., Grazziotin, F.G., Alfaro, M.L., Martins, M., Venzon, M. & Zaher, H. 2016. Diversification in vipers: Phylogenetic relationships, time of divergence and shifts in speciation rates. Molecular Phylogenetics and Evolution 105: 50–62. DOI: 10.1016/j.ympev.2016.07.029 Paywall Reference page.

Vernacular names
беларуская: Ямкагаловыя змеі
čeština: Chřestýšovití
Deutsch: Grubenottern
English: Pit Vipers
hrvatski: Jamičarke
日本語: マムシ亜科
Nederlands: Groefkopadders
polski: Grzechotniki, grzechotnikowate
ไทย: วงศ์ย่อยงูหางกระดิ่ง
Türkçe: Çıngıraklı yılanlar

The Crotalinae, or crotalines, are a subfamily of venomous vipers found in Asia and the Americas. They are distinguished by the presence of a heat-sensing pit organ located between the eye and the nostril on either side of the head. Currently, 18 genera and 151 species are recognized:[4] 7 genera and 54 species in the Old World, against a greater diversity of 11 genera and 97 species in the New World. These are also the only viperids found in the Americas. The groups of snakes represented here include rattlesnakes, lanceheads and Asian pitvipers. The type genus for this subfamily is Crotalus, of which the type species is the timber rattlesnake, C. horridus.

Description

These snakes range in size from the diminutive hump-nosed viper, Hypnale hypnale, that grows to an average only 30–45 centimetres (12–18 in), to the bushmaster, Lachesis muta; a species that is known to reach a maximum of 3.65 metres (12.0 ft) in length -- the longest viperid in the world.
The pit organ is clearly visible between the eye and the nostril of this New Mexico ridge-nosed rattlesnake, Crotalus willardi obscurus.

What makes this group unique is that they all 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 that helps them to find and perhaps even judge the size of the small warm-blooded prey on which they feed.[3]

The pit organ is complex in structure and is similar to the thermoreceptive labial pits found in boas and pythons. It is deep and located in a maxillary cavity. The membrane is like an eardrum that divides the pit into two sections of unequal size, with the larger of the two facing forwards and exposed to the environment. The two sections are connected via a narrow tube, or duct, that can be opened or closed by a group of surrounding muscles. By controlling this tube the snake can balance the air pressure on either side of the membrane.[2] The membrane has many nerve endings packed with mitochondria. Succinic dehydrogenase, lactic dehydrogenase, adenosine triphosphate, monoamine oxidase, generalized esterases and acetylcholine esterase have also been found in it.[3] When prey comes into range, infrared radiation falling onto the membrane allows the snake to determine its direction.[2] Having one of these organs on either side of the head produces a stereo effect that indicates distance as well as direction. Experiments have shown that, when deprived of their senses of sight and smell, these snakes can strike accurately at moving objects that are less than 0.2°C warmer than the background.[5] It would seem as though the pit organs work like a primitive pair of eyes, although it is not known whether the snake experiences this sense as a visual image or in some other fashion.[6] Regardless, it is clear that these organs are of great value to a predator that hunts at night.[7]

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.[3]
Geographic range

This family of snakes is found in the Old World from eastern Europe eastward through Asia to Japan, Taiwan, Indonesia, peninsular India and Sri Lanka. In the Americas, they range from southern Canada southward to Central America to southern South America.[1]
Habitat

This is a versatile group, 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 semi-aquatic: 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.[3]
Behavior

Although a few species are highly active by day, such as Trimeresurus trigonocephalus, a bright green pit viper endemic to Sri Lanka, most are nocturnal, preferring to avoid scorching 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 will 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 that these snakes learn to improve their strike accuracy over time.[8]

Many temperate species (e.g. most rattlesnakes) will congregate in sheltered areas or dens to overwinter (see hibernation), the snakes benefitting from the combined heat. In cool temperatures and while pregnant 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 will strike only if cornered or threatened. Smaller snakes are less likely to stand their ground than larger specimens. Pollution and the destruction of rainforests has caused many viper populations to decline. Humans also threaten vipers, as many vipers are hunted for their skins or killed by cars when they wander onto roads.
Reproduction

With few exceptions, crotalines are ovoviviparous; that is, females give birth to live young. Among the oviparous (egg-laying) pit vipers are Lachesis, Calloselasma, and some Trimeresurus species. It is believed that all egg-laying crotalines guard their eggs.

Brood sizes range from two for very small species, to as many as 86 for the fer-de-lance, Bothrops atrox: a species 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.
Genera

Genus^[4]	Taxon author^[4]	Species^[4]	Subsp.*^[4]	Common name	Geographic range^[1]
Agkistrodon	Palisot de Beauvois, 1799	3	9	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 Yucatan Peninsula, Belize and Guatemala. Along the Pacific coastal plane and lower foothills from Sonora south through Guatemala, El Salvador, Honduras and Nicaragua to northwestern Costa Rica.
Atropoides	Werman, 1992	3	2	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.
Bothriechis	Peters, 1859	7	0	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.
Bothriopsis	Peters, 1861	7	2	Forest-pitvipers	Eastern Panama and most of northern South America, including the Pacific lowlands of Colombia and Ecuador, the Andes Mountains from Venezuela and Colombia to Bolivia, the Amazon Basin and the Atlantic forests of Brazil.
Bothrops	Wagler, 1824	32	11	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	0	Malayan pitviper	Southeast Asia from Thailand to northern Malaysia and Java, Indonesia.
Cerrophidion	Campbell & Lamar, 1992	3	0	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	29	42	Rattlesnakes	The Americas, from southern Canada to northern Argentina.
Deinagkistrodon	Gloyd, 1979	1	0	Hundred-pace pitviper	Southeast Asia.
Gloydius	Hoge & Romano-Hoge, 1981	9	9	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	0	Hump-nosed pit vipers	Sri Lanka and India.
Lachesis	Daudin, 1803	3	1	Bushmasters	Central and South America.
Ophryacus	Cope, 1887	2	0	Mexican horned pitvipers	Mexico.
Ovophis	Burger, 1981	3	4	Mountain pit vipers	Nepal and Seven Sisters (Assam) eastward through Myanmar, Cambodia, Thailand, Laos, Vietnam, West Malaysia, Taiwan, Japan (Okinawa) and Indonesia (Sumatra and Borneo).
Porthidium	Cope, 1871	7	3	Hognose pit vipers	Mexico (Colima, Oaxaca and Chiapas on the Pacific side, the Yucatan Peninsula on the Atlantic side) southward through Central America to northern South America (Ecuador in the Pacific lowlands, northern Venezuela in the Atlantic lowlands).
Sistrurus	Garman, 1883	3	6	Ground rattlesnakes	Southeastern Canada, eastern and northwestern USA, isolated populations in northern and central Mexico.
Trimeresurus	Lacépède, 1804	35	12	Asian lanceheads	Southest Asia from India to southern China and Japan, and the Malay Archipelago to Timor.
Tropidolaemus	Wagler, 1830	2	0	Temple vipers	Southern India and Southeast Asia.

*) Not including the nominate subspecies.
T) Type genus.[1]
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.
See also

* List of crotaline species and subspecies
* Crotalinae by common name
* Crotalinae by taxonomic synonyms
* Snakebite

References

1. ^ a b c d 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).
2. ^ a b c Mehrtens JM. 1987. Living Snakes of the World in Color. New York: Sterling Publishers. 480 pp. ISBN 0-8069-6460-X.
3. ^ a b c d e Campbell JA, Lamar WW. 2004. The Venomous Reptiles of the Western Hemisphere. Comstock Publishing Associates, Ithaca and London. 870 pp. 1500 plates. ISBN 0-8014-4141-2.
4. ^ a b c d e "Crotalinae". Integrated Taxonomic Information System. http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=634394. Retrieved 26 October 2006.
5. ^ Parker HW, Grandison AGC. 1977. Snakes -- a natural history. Second Edition. 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).
6. ^ Bullock, T. H. and Diecke, F. P. J. (1956). Properties of an infrared receptor. Journal of Physiology 134, 47-87.
7. ^ Stidworthy J. 1974. Snakes of the World. Grosset & Dunlap Inc. 160 pp. ISBN 0-448-11856-4.
8. ^ 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 at University of Sydney School of Biological Sciences. Accessed 26 October 2006.