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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...

of vipers found in Asia and the Americas. Like all other vipers, they are venomous. 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, 23

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

and 155 species are recognized: 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 In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...

for this subfamily is '' Crotalus'', of which the

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

is the

timber rattlesnake The timber rattlesnake (''Crotalus horridus''), also known Common name, commonly as the canebrake rattlesnake and the banded rattlesnake,Albert Hazen WWright AH, species:Anna Allen WWright AA (1957). ''Handbook of Snakes of the United States a ...

, ''C. horridus''. These snakes range in size from the diminutive hump-nosed viper, '' Hypnale hypnale'', that grows to a typical total length (including tail) of only , to the bushmaster, '' Lachesis muta'', a species known to reach a maximum total length of in length. This subfamily is unique in 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 Infrared (IR; sometimes called infrared light) is electromagnetic radiation (EMR) with wavelengths longer than that of visible light but shorter than microwaves. The infrared spectral band begins with the waves that are just longer than those ...

-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.Campbell JA, Lamar WW (2004). ''The Venomous Reptiles of the Western Hemisphere''. Ithaca and London: Comstock Publishing Associates. 870 pp., 1,500 plates. . 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. The membrane has many nerve endings packed with

mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...

. Succinic dehydrogenase, lactic dehydrogenase,

adenosine triphosphate Adenosine triphosphate (ATP) is a nucleoside triphosphate that provides energy to drive and support many processes in living cell (biology), cells, such as muscle contraction, nerve impulse propagation, and chemical synthesis. Found in all known ...

monoamine oxidase Monoamine oxidases (MAO) () are a family of enzymes that catalyze the oxidation of monoamines, employing oxygen to clip off their amine group. They are found bound to the outer membrane of mitochondria in most cell types of the body. The fi ...

, generalized esterases, and acetylcholine esterase have also been found in it. When prey comes into range, infrared radiation falling onto the membrane allows the snake to determine its direction. 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, when deprived of their senses of sight and smell, these snakes can strike accurately at moving objects less than warmer than the background.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. (cloth), (paper). The paired pit organs provide the snake with thermal rangefinder capabilities.Bullock, T. H. and Diecke, F. P. J. (1956). Properties of an infrared receptor. Journal of Physiology 134, 47-87. These organs are of great value to a predator that hunts at night, as well as for avoiding the snake’s own predators.Stidworthy J. 1974. Snakes of the World. Grosset & Dunlap Inc. 160 pp. . 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 muscularis compressor glandulae, forces venom out of the gland.

Evolution

The earliest known fossil pit viper remains are from the

Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...

Nebraska Nebraska ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Ka ...

. As pit vipers are thought to have had an Asian origin before eventually colonizing the Americas, this suggests that they must have originated and diversified even earlier. During the Late Miocene, they reached as far west as eastern Europe, where they are no longer found; it is thought that they did not expand further into Europe.

Geographic range

The subfamily Crotalinae is found from

Central Asia Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...

eastward and southward to

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...

, peninsular

India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

Nepal Nepal, officially the Federal Democratic Republic of Nepal, is a landlocked country in South Asia. It is mainly situated in the Himalayas, but also includes parts of the Indo-Gangetic Plain. It borders the Tibet Autonomous Region of China Ch ...

Bangladesh Bangladesh, officially the People's Republic of Bangladesh, is a country in South Asia. It is the List of countries and dependencies by population, eighth-most populous country in the world and among the List of countries and dependencies by ...

and

Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...

. In the Americas, they range from southern

Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...

southward to Central America to southern South America.

Habitat

Crotalines are a versatile subfamily, with members found in

habitats In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

ranging from parched

desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...

(e.g., the sidewinder, '' Crotalus cerastes'') to

rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...

s (e.g., the bushmaster, ''Lachesis muta''). They may be either

arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...

or terrestrial, and at least one species (the cottonmouth, '' Agkistrodon piscivorus'') is semiaquatic. 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 above sea level.

Behavior

Although a few species of crotalines are highly active by day, such as '' Trimeresurus trigonocephalus'', a bright green pit viper

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

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. 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 Sydne
School of Biological Sciences
Accessed 26 October 2006. Many temperate species of pit vipers (e.g. most rattlesnakes) congregate in sheltered areas or "dens" to overwinter (brumate, 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, when 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 Oviparous animals are animals that reproduce by depositing fertilized zygotes outside the body (i.e., by laying or spawning) in metabolically independent incubation organs known as eggs, which nurture the embryo into moving offsprings kno ...

(egg-laying) pit vipers are '' Lachesis'', '' Calloselasma'', and some ''

Trimeresurus ''Trimeresurus'' is a genus of pit vipers native to Asia. They are found from the Indian Subcontinent throughout Southeast Asia, China, and the Pacific Islands. The genus currently contains 44 recognized species. Common names include Asian palm p ...

'' 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. These tails are known to be used by a number of species 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 Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

: the Crotalidae. Today, however, the

monophyly In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...

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

''*) Not including the nominate subspecies.''
)

Type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...

References

External links

Pit organs at Life is Short, but Snakes are Long
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