Scorpion: Difference between revisions

Scorpions; Temporal range: 435–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Early Silurian – present
	Hottentotta tamulus from Mangaon, Maharashtra, India
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Chelicerata
Class:	Arachnida
Clade:	Arachnopulmonata
Order:	Scorpiones; C. L. Koch, 1837
Families
	see Taxonomy
	Native range of Scorpiones

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Scorpions are predatory arachnids of the order Scorpiones. They have eight legs and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always ending with a stinger. The evolutionary history of scorpions goes back 435 million years. They mainly live in deserts but have adapted to a wide range of environmental conditions, and can be found on all continents except Antarctica. There are over 2,500 described species, with 22 extant (living) families recognized to date. Their taxonomy is being revised to account for 21st-century genomic studies.

Scorpions primarily prey on insects and other invertebrates, but some species hunt vertebrates. They use their pincers to restrain and kill prey, or to prevent their own predation. The venomous sting is used for offense and defense. During courtship, the male and female grasp each other's pincers and dance while he tries to move her onto his sperm packet. All known species give live birth and the female cares for the young as their exoskeletons harden, transporting them on her back. The exoskeleton contains fluorescent chemicals and glows under ultraviolet light.

The vast majority of species do not seriously threaten humans, and healthy adults usually do not need medical treatment after a sting. About 25 species (fewer than one percent) have venom capable of killing a human, which happens frequently in the parts of the world where they live, primarily where access to medical treatment is unlikely.

Scorpions appear in art, folklore, mythology, and commercial brands. Scorpion motifs are woven into kilim carpets for protection from their sting. Scorpius is the name of a constellation; the corresponding astrological sign is Scorpio. A classical myth about Scorpius tells how the giant scorpion and its enemy Orion became constellations on opposite sides of the sky.

Etymology

The word scorpion originated in Middle English between 1175 and 1225 AD from Old French scorpion,^[1] or from Italian scorpione, both derived from the Latin scorpio, equivalent to scorpius,^[2] which is the romanization of the Greek σκορπίος – skorpíos,^[3] with no native IE etymology (cfr. Arabic ʕaqrab 'scorpion', Proto-Germanic *krabbô 'crab').

Evolution

Fossil record

Scorpion fossils have been found in many strata, including marine Silurian and estuarine Devonian deposits, coal deposits from the Carboniferous Period and in amber. Whether the early scorpions were marine or terrestrial has been debated, though they had book lungs like modern terrestrial species.^[5]^[6]^[7]^[8] Over 100 fossil species of scorpion have been described.^[9] The oldest found as of 2021 is Dolichophonus loudonensis, which lived during the Silurian, in present-day Scotland.^[10] Gondwanascorpio from the Devonian is among the earliest-known terrestrial animals on the Gondwana supercontinent.^[11] Some Palaeozoic scorpions possessed compound eyes similar to those of eurypterids.^[12] The Triassic fossils Protochactas and Protobuthus belong to the modern clades Chactoidea and Buthoidea respectively, indicating that the crown group of modern scorpions had emerged by this time.^[13]

Phylogeny

The Scorpiones are a clade within the pulmonate Arachnida (those with book lungs). Arachnida is placed within the Chelicerata, a subphylum of Arthropoda that contains sea spiders and horseshoe crabs, alongside terrestrial animals without book lungs such as ticks and harvestmen.^[5] The extinct Eurypterida, sometimes called sea scorpions, though they were not all marine, are not scorpions; their grasping pincers were chelicerae, not homologous with the pincers (second appendages) of scorpions.^[14] Scorpiones is sister to the Tetrapulmonata, a terrestrial group of pulmonates containing the spiders and whip scorpions. This 2019 cladogram summarizes:^[5]

Chelicerata

Pycnogonida (sea spiders)

Prosomapoda

Xiphosura (horseshoe crabs)

†Eurypterida (sea scorpions)

Arachnida

Non‑pulmonates

(ticks, harvestmen, etc)

pulmonates

Scorpiones

Tetrapulmonata

	Araneae (spiders)

	Pedipalpi (whip scorpions, etc)

Recent studies place pseudoscorpions as the sister group of scorpions in the clade Panscorpiones, which together with Tetrapulmonata makes up the clade Arachnopulmonata.^[15]

The internal phylogeny of the scorpions has been debated,^[5] but genomic analysis consistently places the Bothriuridae as sister to a clade consisting of Scorpionoidea and Chactoidea. The scorpions diversified between the Devonian and the early Carboniferous. The main division is into the clades Buthida and Iurida. The Bothriuridae diverged starting before temperate Gondwana broke up into separate land masses, completed by the Jurassic. The Iuroidea and Chactoidea are both seen not to be single clades, and are shown as "paraphyletic" (with quotation marks) in this 2018 cladogram.^[16]

Scorpiones

Buthida

	Chaeriloidea

	Pseudochactoidea

Buthoidea

Iurida

"Iuroidea" (part)

Bothriuroidea

	"Chactoidea" (part)

	"Iuroidea" (part)

"Chactoidea" (part)

Scorpionoidea

Taxonomy

Carl Linnaeus described six species of scorpion in his genus Scorpio in 1758 and 1767; three of these are now considered valid and are called Scorpio maurus, Androctonus australis, and Euscorpius carpathicus; the other three are dubious names. He placed the scorpions among his "Insecta aptera" (wingless insects), a group that included Crustacea, Arachnida and Myriapoda.^[17] In 1801, Jean-Baptiste Lamarck divided up the "Insecta aptera", creating the taxon Arachnides for spiders, scorpions, and acari (mites and ticks), though it also contained the Thysanura, Myriapoda and parasites such as lice.^[18] German arachnologist Carl Ludwig Koch created the order Scorpiones in 1837. He divided it into four families, the six-eyed scorpions "Scorpionides", the eight-eyed scorpions "Buthides", the ten-eyed scorpions "Centrurides", and the twelve-eyed scorpions "Androctonides".^[19]

More recently, some twenty-two families containing over 2,500 species of scorpions have been described, with many additions and much reorganization of taxa in the 21st century.^[20]^[5]^[21] There are over 100 described taxa of fossil scorpions.^[9] This classification is based on Soleglad and Fet (2003),^[22] which replaced Stockwell's older, unpublished classification.^[23] Further taxonomic changes are from papers by Soleglad et al. (2005).^[24]^[25]

The extant taxa to the rank of family (numbers of species in parentheses^[20]) are:

Order Scorpiones

*Centruroides vittatus*, the striped bark scorpion, a member of Buthidae, the largest family of scorpions

Parvorder Pseudochactida Soleglad & Fet, 2003
- Superfamily Pseudochactoidea Gromov, 1998
  - Family Pseudochactidae Gromov, 1998 (1 sp.) (Central Asian scorpions of semi-savanna habitats)
Parvorder Buthida Soleglad & Fet, 2003
- Superfamily Buthoidea C. L. Koch, 1837
  - Family Buthidae C. L. Koch, 1837 (1209 spp.) (thick-tailed scorpions, including the most dangerous species)
  - Family Microcharmidae Lourenço, 1996, 2019 (17 spp.) (African scorpions of humid forest leaf litter)
Parvorder Chaerilida Soleglad & Fet, 2003
- Superfamily Chaeriloidea Pocock, 1893
  - Family Chaerilidae Pocock, 1893 (51 spp.) (South and Southeast Asian scorpions of non-arid places)

Parvorder Iurida Soleglad & Fet, 2003
- Superfamily Chactoidea Pocock, 1893
  - Family Akravidae Levy, 2007 (1 sp.) (cave-dwelling scorpions of Israel)
  - Family Belisariidae Lourenço, 1998 (3 spp.) (cave-related scorpions of Southern Europe)
  - Family Chactidae Pocock, 1893 (209 spp.) (New World scorpions, membership under revision)
  - Family Euscorpiidae Laurie, 1896 (170 spp.) (harmless scorpions of the Americas, Eurasia, and North Africa)
  - Family Superstitioniidae Stahnke, 1940 (1 sp.) (cave scorpions of Mexico and Southwestern United States)
  - Family Troglotayosicidae Lourenço, 1998 (4 spp.) (cave-related scorpions of South America)
  - Family Typhlochactidae Mitchell, 1971 (11 spp.) (cave-related scorpions of Eastern Mexico)
  - Family Vaejovidae Thorell, 1876 (222 spp.) (New World scorpions)
- Superfamily Iuroidea Thorell, 1876
  - Family Caraboctonidae Kraepelin, 1905 (23 spp.) (hairy scorpions)
  - Family Hadruridae Stahnke, 1974 (9 spp.) (large North American scorpions)
  - Family Iuridae Thorell, 1876 (21 spp.) (scorpions with a large tooth on inner side of moveable claw)
- Superfamily Scorpionoidea Latreille, 1802
  - Family Bothriuridae Simon, 1880 (158 spp.) (Southern hemisphere tropical and temperate scorpions)
  - Family Hemiscorpiidae Pocock, 1893 (16 spp.) (rock, creeping, or tree scorpions of the Middle East)
  - Family Hormuridae Laurie, 1896 (92 spp.) (flattened, crevice-living scorpions of Southeast Asia and Australia)
  - Family Rugodentidae Bastawade et al., 2005 (1 sp.) (burrowing scorpions of India)
  - Family Scorpionidae Latreille, 1802 (183 spp.) (burrowing or pale-legged scorpions)
  - Family Diplocentridae Karsch, 1880 (134 spp.) (closely related to and sometimes placed in Scorpionidae, but have spine on telson)
  - Family Heteroscorpionidae Kraepelin, 1905 (6 spp.) (scorpions of Madagascar)

Geographical distribution

Scorpions are found on all continents except Antarctica. The diversity of scorpions is greatest in subtropical areas; it decreases toward the poles and equator, though scorpions are found in the tropics. Scorpions did not occur naturally in Great Britain but were accidentally introduced by humans, and have now established a population.^[26]^[27] New Zealand, and some of the islands in Oceania, have in the past had small populations of introduced scorpions, but they were exterminated.^[28]^[26] Five colonies of Euscorpius flavicaudis have established themselves since the late 19th century in Sheerness in England at 51°N,^[29]^[30]^[31] while Paruroctonus boreus lives as far north as Red Deer, Alberta, at 52°N.^[32] A few species are on the IUCN Red List; Lychas braueri is classed as critically endangered (2014), Isometrus deharvengi as endangered (2016) and Chiromachus ochropus as vulnerable (2014).^[33]^[34]^[35]

Scorpions are xerocoles, meaning they primarily live in deserts, but they can be found in virtually every terrestrial habitat including high-elevation mountains, caves, and intertidal zones. They are largely absent from boreal ecosystems such as the tundra, high-altitude taiga, and mountain tops.^[36]^[5] The highest altitude reached by a scorpion is 5,500 meters (18,000 ft) in the Andes, for Orobothriurus crassimanus.^[37]

As regards microhabitats, scorpions may be ground-dwelling, tree-loving, rock-loving or sand-loving. Some species, such as Vaejovis janssi, are versatile and are found in all habitats on Socorro Island, Baja California, while others such as Euscorpius carpathicus, endemic to the littoral zone of rivers in Romania, occupy specialized niches.^[38]^[39]

Morphology

Scorpions range in size from the 8.5 mm (0.33 in) Typhlochactas mitchelli of Typhlochactidae,^[38] to the 23 cm (9.1 in) Heterometrus swammerdami of Scorpionidae.^[40] The body of a scorpion is divided into two parts or tagmata: the cephalothorax or prosoma, and the abdomen or opisthosoma.^[a] The opisthosoma is subdivided into a broad anterior portion, the mesosoma or pre-abdomen, and a narrow tail-like posterior, the metasoma or post-abdomen.^[42] External differences between the sexes are not obvious in most species. In some, the metasoma is more elongated in males than females.^[43]

Cephalothorax

The cephalothorax comprises the carapace, eyes, chelicerae (mouth parts), pedipalps (which have chelae, commonly called claws or pincers) and four pairs of walking legs. Scorpions have two eyes on the top of the cephalothorax, and usually two to five pairs of eyes along the front corners of the cephalothorax. While unable to form sharp images, their central eyes are amongst the most light sensitive in the animal kingdom, especially in dim light, which makes it possible for nocturnal species to use starlight to navigate at night.^[44] The chelicerae are at the front and underneath the carapace. They are pincer-like and have three segments and sharp "teeth".^[45]^[46] The brain of a scorpion is in the back of the cephalothorax, just above the esophagus.^[47] As in other arachnids, the nervous system is highly concentrated in the cephalothorax, but has a long ventral nerve cord with segmented ganglia which may be a primitive trait.^[48]

The pedipalp is a segmented, clawed appendage used for prey immobilization, defense and sensory purposes. The segments of the pedipalp (from closest to the body outward) are coxa, trochanter, femur, patella, tibia (including the fixed claw and the manus) and tarsus (moveable claw). A scorpion has darkened or granular raised linear ridges, called "keels" or "carinae" on the pedipalp segments and on other parts of the body; these are useful as taxonomic characters.^[49] Unlike those of some other arachnids, the legs have not been modified for other purposes, though they may occasionally be used for digging, and females may use them to catch emerging young. The legs are covered in proprioceptors, bristles and sensory setae.^[50] Depending on the species, the legs may have spines and spurs.^[51]

Mesosoma

The mesosoma or preabdomen is the broad part of the opisthosoma.^[42] In the early stages of embryonic development the mesosoma consist of eight segments, but the first segment disappear before birth, so the mesosoma in scorpions actually consist of segments 2-8.^[52]^[53]^[54] These anterior seven somites (segments) of the opisthosoma are each covered dorsally by a sclerotized plate called the tergite. Ventrally, somites 3 to 7 are armored with matching plates called sternites. The ventral side of somite 1 has a pair of genital opercula covering the gonopore. Sternite 2 forms the basal plate bearing the pectines,^[55] which function as sensory organs.^[56]

The next four somites, 3 to 6, all bear pairs of spiracles. They serve as openings for the scorpion's respiratory organs, known as book lungs. The spiracle openings may be slits, circular, elliptical or oval according to the species.^[57]^[58] There are thus four pairs of book lungs; each consists of some 140 to 150 thin lamellae filled with air inside a pulmonary chamber, connected on the ventral side to an atrial chamber which opens into a spiracle. Bristles hold the lamellae apart. A muscle opens the spiracle and widens the atrial chamber; dorsoventral muscles contract to compress the pulmonary chamber, forcing air out, and relax to allow the chamber to refill.^[59] The 7th and last somite does not bear appendages or any other significant external structures.^[57]

The mesosoma contains the heart or "dorsal vessel" which is the center of the scorpion's open circulatory system. The heart is continuous with a deep arterial system which spreads throughout the body. Sinuses return deoxygenated blood (hemolymph) to the heart; the blood is re-oxygenated by cardiac pores. The mesosoma also contains the reproductive system. The female gonads are made of three or four tubes that run parallel to each other and are connected by two to four transverse anastomoses. These tubes are the sites for both oocyte formation and embryonic development. They connect to two oviducts which connect to a single atrium leading to the genital orifice.^[60] Males have two gonads made of two cylindrical tubes with a ladder-like configuration; they contain cysts which produce spermatozoa. Both tubes end in a spermiduct, one on each side of the mesosoma. They connect to glandular symmetrical structures called paraxial organs, which end at the genital orifice. These secrete chitin-based structures which come together to form the spermatophore.^[61]^[62]

Metasoma

The "tail" or metasoma consists of five segments and the telson, which is not strictly a segment. The five segments are merely body rings; they lack apparent sterna or terga, and become larger distally. These segments have keels, setae and bristles which may be used for taxonomic classification. The anus is at the distal and ventral end of the last segment, and is encircled by four anal papillae and the anal arch.^[57] The tails of some species contain light receptors.^[44]

The telson includes the vesicle, which contains a symmetrical pair of venom glands. Externally it bears the curved stinger, the hypodermic aculeus, equipped with sensory hairs. Each of the venom glands has its own duct to convey its secretion along the aculeus from the bulb of the gland to immediately near of the tip, where each of the paired ducts has its own venom pore.^[63] An extrinsic muscle system in the tail moves it forward and propels and penetrates with the aculeus, while an intrinsic muscle system attached to the glands pumps venom through the stinger into the intended victim.^[64] The stinger contains metalloproteins with zinc, hardening the tip.^[65] The optimal stinging angle is around 30 degrees relative to the tip.^[66]

Biology

Most scorpion species are nocturnal or crepuscular, finding shelter during the day in burrows, cracks in rocks and tree bark.^[67] Many species dig a shelter underneath stones a few centimeters long. Some may use burrows made by other animals including spiders, reptiles and small mammals. Other species dig their own burrows which vary in complexity and depth. Hadrurus species dig burrows as over 2 m (6 ft 7 in) deep. Digging is done using the mouth parts, claws and legs. In several species, particularly of the family Buthidae, individuals may gather in the same shelter; bark scorpions may aggregate up to 30 individuals. In some species, families of females and young sometimes aggregate.^[68]

Scorpions prefer areas where the temperature remains in the range of 11–40 °C (52–104 °F), but may survive temperatures from well below freezing to desert heat.^[69]^[70] Scorpions can withstand intense heat: Leiurus quinquestriatus, Scorpio maurus and Hadrurus arizonensis can live in temperatures of 45–50 °C (113–122 °F) if they are sufficiently hydrated. Desert species must deal with the extreme changes in temperature from day to night or between seasons; Pectinibuthus birulai lives in a temperature range of −30–50 °C (−22–122 °F). Scorpions that live outside deserts prefer lower temperatures. The ability to resist cold may be related to the increase in the sugar trehalose when the temperature drops. Some species hibernate.^[71] Scorpions appear to have resistance to ionizing radiation. This was discovered in the early 1960s when scorpions were found to be among the few animals to survive nuclear tests at Reggane, Algeria.^[72]

Desert scorpions have several adaptations for water conservation. They excrete insoluble compounds such as xanthine, guanine, and uric acid, not requiring water for their removal from the body. Guanine is the main component and maximizes the amount of nitrogen excreted. A scorpion's cuticle holds in moisture via lipids and waxes from epidermal glands, and protects against ultraviolet radiation. Even when dehydrated, a scorpion can tolerate high osmotic pressure in its blood.^[73] Desert scorpions get most of their moisture from the food they eat but some can absorb water from the humid soil. Species that live in denser vegetation and in more moderate temperatures will drink water on plants and in puddles.^[74]

A scorpion uses its stinger both for killing prey and defense. Some species make direct, quick strikes with their tails while others make slower, more circular strikes which can more easily return the stinger to a position where it can strike again. Leiurus quinquestriatus can whip its tail at a speed of up to 128 cm/s (50 in/s) in a defensive strike.^[75]

Mortality and defense

Scorpions may be attacked by other arthropods like ants, spiders, solifugids and centipedes. Major predators include frogs, lizards, snakes, birds, and mammals.^[76] Meerkats are somewhat specialized in preying on scorpions, biting off their stingers and being immune to their venom.^[77]^[78] Other predators adapted for hunting scorpions include the grasshopper mouse and desert long-eared bat, which are also immune to their venom.^[79]^[80] In one study, 70% of the latter's droppings contained scorpion fragments.^[80] Scorpions host parasites including mites, scuttle flies, nematodes and some bacteria. The immune system of scorpions gives them resistance to infection by many types of bacteria.^[81]

When threatened, a scorpion raises its claws and tail in a defensive posture. Some species stridulate to warn off predators by rubbing certain hairs, the stinger or the claws.^[76] Certain species have a preference for using either the claws or stinger as defense, depending on the size of the appendages.^[82] A few scorpions, such as Parabuthus, Centruroides margaritatus, and Hadrurus arizonensis, squirt venom in a narrow jet as far as 1 meter (3.3 ft) to warn off potential predators, possibly injuring them in the eyes.^[83] Some Ananteris species can shed parts of their tail to escape predators. The parts do not grow back, leaving them unable to sting and defecate, but they can still catch small prey and reproduce for at least eight months afterward.^[84]

Diet and feeding

Scorpions generally prey on insects, particularly grasshoppers, crickets, termites, beetles and wasps. Other prey include spiders, solifugids, woodlice and even small vertebrates including lizards, snakes and mammals. Species with large claws may prey on earthworms and mollusks. The majority of species are opportunistic and consume a variety of prey though some may be highly specialized; Isometroides vescus specializes on burrowing spiders. Prey size depends on the size of the species. Several scorpion species are sit-and-wait predators, which involves them waiting for prey at or near the entrance to their burrow. Others actively seek them out. Scorpions detect their prey with mechanoreceptive and chemoreceptive hairs on their bodies and capture them with their claws. Small animals are merely killed with the claws, particularly by large-clawed species. Larger and more aggressive prey is given a sting.^[85]^[86]

Scorpions, like other arachnids, digest their food externally. The chelicerae, which are very sharp, are used to pull small amounts of food off the prey item into a pre-oral cavity below the chelicerae and carapace. The digestive juices from the gut are egested onto the food, and the digested food is then sucked into the gut in liquid form. Any solid indigestible matter (such as exoskeleton fragments) is trapped by setae in the pre-oral cavity and ejected. The sucked-in food is pumped into the midgut by the pharynx, where it is further digested. The waste passes through the hindgut and out of the anus. Scorpions can consume large amounts of food during one meal. They have an efficient food storage organ and a very low metabolic rate, and a relatively inactive lifestyle. This enables some to survive six to twelve months of starvation.^[87]

Mating

Most scorpions reproduce sexually, with male and female individuals; species in some genera, such as Hottentotta and Tityus, and the species Centruroides gracilis, Liocheles australasiae, and Ananteris coineaui have been reported, not necessarily reliably, to reproduce through parthenogenesis, in which unfertilized eggs develop into living embryos.^[88] Receptive females produce pheromones which are picked up by wandering males using their pectines to comb the substrate. Males begin courtship by moving their bodies back and forth, without moving the legs, a behavior known as juddering. This appears to produce ground vibrations that are picked up by the female.^[61]

The pair then make contact using their pedipalps, and perform a dance called the promenade à deux (French for "a walk for two"). In this dance, the male and female move back and forth while facing each other, as the male searches for a suitable place to deposit his spermatophore. The courtship ritual can involve several other behaviors such as a cheliceral kiss, in which the male and female grasp each other's mouth-parts, arbre droit ("upright tree") where the partners elevate their posteriors and rub their tails together, and sexual stinging, in which the male stings the female in the chelae or mesosoma to subdue her. The dance can last from a few minutes to several hours.^[89]^[90]

When the male has located a suitably stable substrate, such as hard ground, agglomerated sand, rock, or tree bark, he deposits the spermatophore and guides the female over it. This allows the spermatophore to enter her genital opercula, which triggers release of the sperm, thus fertilizing the female. A mating plug then forms in the female to prevent her from mating again before the young are born. The male and female then abruptly separate.^[91]^[92] Sexual cannibalism after mating has only been reported anecdotally in scorpions.^[93]

Birth and development

*Compsobuthus werneri* female with young

Gestation in scorpions can last for over a year in some species.^[94] They have two types of embryonic development; apoikogenic and katoikogenic. In the apoikogenic system, which is mainly found in the Buthidae, embryos develop in yolk-rich eggs inside follicles. The katoikogenic system is documented in Hemiscorpiidae, Scorpionidae and Diplocentridae, and involves the embryos developing in a diverticulum which has a teat-like structure for them to feed through.^[95] Unlike the majority of arachnids, which are oviparous, hatching from eggs, scorpions seem to be universally viviparous, with live births.^[96] They are unusual among terrestrial arthropods in the amount of care a female gives to her offspring.^[97] The size of a brood varies by species, from 3 to over 100.^[98] The body size of scorpions is not correlated either with brood size or with life cycle length.^[99]

Before giving birth, the female elevates the front of her body and positions her pedipalps and front legs under her to catch the young ("birth basket"). The young emerge one by one from the genital opercula, expel the embryonic membrane, if any, and are placed on the mother's back where they remain until they have gone through at least one molt. The period before the first molt is called the pro-juvenile stage; the young are unable to feed or sting, but have suckers on their tarsi, used to hold on to their mother. This period lasts 5 to 25 days, depending on the species. The brood molt for the first time simultaneously in a process that lasts 6 to 8 hours, marking the beginning of the juvenile stage.^[98]

Juvenile stages or instars generally resemble smaller versions of adults, with fully developed pincers, hairs and stingers. They are still soft and lack pigments, and thus continue to ride on their mother's back for protection. They become harder and more pigmented over the next couple of days. They may leave their mother temporarily, returning when they sense potential danger. Once the exoskeleton is fully hardened, the young can hunt prey on their own and may soon leave their mother.^[100] A scorpion may molt six times on average before reaching maturity, which may not occur until it is 6 to 83 months old, depending on the species. Some species may live up to 25 years.^[94]

Fluorescence

This black emperor scorpion (Pandinus imperator) fluoresces light blue.

The mother glows bright teal, the young a dull grey.

Scorpions glow a vibrant blue-green when exposed to certain wavelengths of ultraviolet light, such as that produced by a black light, due to fluorescent chemicals such as beta-carboline in the cuticle. Accordingly, a hand-held ultraviolet lamp has long been a standard tool for nocturnal field surveys of these animals. Fluorescence occurs as a result of sclerotization and increases in intensity with each successive instar.^[101] This fluorescence may have an active role in the scorpion's ability to detect light.^[102]

Relationship with humans

Stings

Arizona bark scorpion, one of the few species whose venom is deadly to humans

Scorpion venom serves to kill or paralyze prey rapidly. The stings of many species are uncomfortable, but only 25 species have venom that is deadly to humans. Those species belong to the family Buthidae, including Leiurus quinquestriatus, Hottentotta spp., Centruroides spp., and Androctonus spp.^[38] People with allergies are especially at risk;^[103] otherwise, first aid is symptomatic, with analgesia. Cases of very high blood pressure are treated with medications that relieve anxiety and relax the blood vessels.^[104]^[105] Scorpion envenomation with high morbidity and mortality is usually due to either excessive autonomic activity and cardiovascular toxic effects, or neuromuscular toxic effects. Antivenom is the specific treatment for scorpion envenomation combined with supportive measures including vasodilators in patients with cardiovascular toxic effects, and benzodiazepines when there is neuromuscular involvement. Although rare, severe hypersensitivity reactions including anaphylaxis to scorpion antivenin are possible.^[106]

Scorpion stings are a public health problem, particularly in the tropical and subtropical regions of the Americas, North Africa, the Middle East and India. Around 1.5 million scorpion envenomations occur each year with around 2,600 deaths.^[107]^[108]^[109] Mexico is one of the most affected countries, with the highest biodiversity of scorpions in the world, some 200,000 envenomations per year and at least 300 deaths.^[110]^[111]

Efforts are made to prevent envenomation and to control scorpion populations. Prevention encompasses personal activities such as checking shoes and clothes before putting them on, not walking in bare feet or sandals, and filling in holes and cracks where scorpions might nest. Street lighting reduces scorpion activity. Control may involve the use of insecticides such as pyrethroids, or gathering scorpions manually with the help of ultraviolet lights. Domestic predators of scorpions, such as chickens and turkeys, can help to reduce the risk to a household.^[107]^[108]

Potential medicinal use

The deathstalker's powerful venom contains the 36-amino acid peptide chlorotoxin (ribbon diagram shown). This blocks small-conductance chloride channels, immobilizing its prey.^[112]

Scorpion venom is a mixture of neurotoxins; most of these are peptides, chains of amino acids.^[113] Many of them interfere with membrane channels that transport sodium, potassium, calcium, or chloride ions. These channels are essential for nerve conduction, muscle contraction and many other biological processes. Some of these molecules may be useful in medical research and might lead to the development of new disease treatments. Among their potential therapeutic uses are as analgesic, anti-cancer, antibacterial, antifungal, antiviral, antiparasitic, bradykinin-potentiating, and immunosuppressive drugs. As of 2020, no scorpion toxin-based drug is for sale, though chlorotoxin is being trialled for use against glioma, a brain cancer.^[112]

Consumption

Scorpions are eaten by people in West Africa, Myanmar^[114] and East Asia. Fried scorpion is traditionally eaten in Shandong, China.^[115] There, scorpions can be cooked and eaten in a variety of ways, including roasting, frying, grilling, raw, or alive. The stingers are typically not removed, since direct and sustained heat negates the harmful effects of the venom.^[116] In Thailand, scorpions are not eaten as often as other arthropods, such as grasshoppers, but they are sometimes fried as street food.^[117] They are used in Vietnam to make snake wine (scorpion wine).^[118]

Pets

Scorpions are often kept as pets. They are relatively simple to keep, the main requirements being a secure enclosure such as a glass terrarium with a lockable lid and the appropriate temperature and humidity for the chosen species, which typically means installing a heating mat and spraying regularly with a little water. The substrate needs to resemble that of the species' natural environment, such as peat for forest species, or lateritic sand for burrowing desert species. Scorpions in the genera Pandinus and Heterometrus are docile enough to handle. A large Pandinus may consume up to three crickets each week. Cannibalism is more common in captivity than in the wild and can be minimized by providing many small shelters within the enclosure and ensuring there is plenty of prey.^[119]^[120] The pet trade has threatened wild populations of some scorpion species, particularly Androctonus australis and Pandinus imperator.^[121]

Culture

Late period bronze figure of Isis-Serket
"Scorpion and snake fighting", Anglo-Saxon Herbal, c. 1050
The constellation Scorpius, depicted in Urania's Mirror as "Scorpio", London, c. 1825
A scorpion motif (two types shown) was often woven into Turkish kilim flatweave carpets, for protection from their sting.^[122]

The scorpion is a culturally significant animal, appearing as a motif in art, especially in Islamic art in the Middle East.^[123] A scorpion motif is often woven into Turkish kilim flatweave carpets, for protection from their sting.^[122] The scorpion is perceived both as an embodiment of evil and as a protective force such as a dervish's powers to combat evil.^[123] In Muslim folklore, the scorpion portrays human sexuality.^[123] Scorpions are used in folk medicine in South Asia, especially in antidotes for scorpion stings.^[123]

One of the earliest occurrences of the scorpion in culture is its inclusion, as Scorpio, in the 12 signs of the Zodiac by Babylonian astronomers during the Chaldean period. This was then taken up by western astrology; in astronomy the corresponding constellation is named Scorpius.^[124] In ancient Egypt, the goddess Serket, who protected the Pharaoh, was often depicted as a scorpion.^[125] In ancient Greece, a warrior's shield sometimes carried a scorpion device, as seen in red-figure pottery from the 5th century BC.^[126] In Greek mythology, Artemis or Gaia sent a giant scorpion to kill the hunter Orion, who had said he would kill all the world's animals. Orion and the scorpion both became constellations; as enemies they were placed on opposite sides of the world, so when one rises in the sky, the other sets.^[127]^[128] Scorpions are mentioned in the Bible and the Talmud as symbols of danger and maliciousness.^[128]

The fable of The Scorpion and the Frog has been interpreted as showing that vicious people cannot resist hurting others, even when it is not in their interests.^[129] More recently, the action in John Steinbeck's 1947 novella The Pearl centers on a poor pearl fisherman's attempts to save his infant son from a scorpion sting, only to lose him to human violence.^[130] Scorpions have equally appeared in western artforms including film and poetry: the surrealist filmmaker Luis Buñuel made symbolic use of scorpions in his 1930 classic L'Age d'or (The Golden Age),^[131] while Stevie Smith's last collection of poems was entitled Scorpion and other Poems.^[132] A variety of martial arts films and video games have been entitled Scorpion King.^[133]^[134]^[135]

Scorpion pose in yoga has one or both legs pointing forward over the head, like a scorpion's tail.^[136]

Since classical times, the scorpion with its powerful stinger has been used to provide a name for weapons. In the Roman army, the scorpio was a torsion siege engine used to shoot a projectile.^[137] The British Army's FV101 Scorpion was an armored reconnaissance vehicle or light tank in service from 1972 to 1994.^[138] A version of the Matilda II tank, fitted with a flail to clear mines, was named the Matilda Scorpion.^[139] Several ships of the Royal Navy and of the US Navy have been named Scorpion including an 18-gun sloop in 1803,^[140] a turret ship in 1863,^[141] a patrol yacht in 1898,^[142] a destroyer in 1910,^[143] and a nuclear submarine in 1960.^[144]

The scorpion has served as the name or symbol of products and brands including Italy's Abarth racing cars^[145] and a Montesa scrambler motorcycle.^[146] A hand- or forearm-balancing asana in modern yoga as exercise with the back arched and one or both legs pointing forward over the head in the manner of the scorpion's tail is called Scorpion pose.^[147]^[136]

Notes

^ As there is currently neither paleontological nor embryological evidence that arachnids ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all scorpions contains a heart and book lungs, organs atypical of an abdomen.^[41]

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External links

[42] As there is currently neither paleontological nor embryological evidence that arachnids ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all scorpions contains a heart and book lungs, organs atypical of an abdomen.^[41]

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[127] ttrib. to "The Achilles Painter". "Pelike". British Museum. Archived from the original on 3 October 2020. Retrieved 24 August 2020. in his left [hand] he carries a long spear and a shield with device in black silhouette of a scorpion to left. ... Made in: Attica, Greece ... Findspot: Nola, Italy

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@@ Line 1: / Line 1: @@
 {{short description|Predatory order of arachnids}}
 {{About |the arthropod}}
+{{hatnote|"Scorpion tail" and "scorpion's tail" redirect here. For other uses, see [[Scorpion tail (disambiguation)]].}}
+{{pp-semi-indef}}
 {{Use dmy dates|date=May 2021}}
 {{Automatic taxobox
  | name              = Scorpions
+ | display_parents   = 2
  | taxon             = Scorpiones
- | image             = Scorpiones-01.jpg
+ | image = Scorpion_Photograph_By_Shantanu_Kuveskar.jpg
- | image_caption     =  Scorpiones
+ | image_caption = ''[[Hottentotta tamulus]]'' from [[Mangaon]], [[Maharashtra]], [[India]]
- | fossil_range      = {{Fossil range |435|0}}<small>[[Llandovery epoch|Early Silurian]] – [[Holocene|present]]</small>
+ | fossil_range      = {{Fossil range |435|0}}<small>[[Early Silurian]] – [[Holocene|present]]</small>
  | authority         = [[C. L. Koch]], 1837
  | subdivision_ranks = Families
@@ Line 14: / Line 17: @@
 | range_map_caption  = Native range of Scorpiones
 }}
-'''Scorpions''' are [[predatory]] [[arachnid]]s of the [[Order (biology)|order]] '''Scorpiones'''. They have eight legs, and are easily recognized by a pair of [[Chela (organ)|grasping pincers]] and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always ending with a [[stinger]]. The evolutionary history of scorpions goes back [[Silurian|435 million years]]. They mainly live in [[desert]]s but have adapted to a wide range of environmental conditions, and can be found on all continents except [[Antarctica]]. There are over 2,500 described [[species]], with 22 extant (living) families recognized to date. Their [[Taxonomy (biology)|taxonomy]] is being revised to account for 21st-century [[genomic]] studies.
+'''Scorpions''' are [[predatory]] [[arachnid]]s of the [[Order (biology)|order]] '''Scorpiones'''. They have eight legs and are easily recognized by a pair of [[Chela (organ)|grasping pincers]] and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always ending with a [[stinger]]. The evolutionary history of scorpions goes back [[Silurian|435 million years]]. They mainly live in [[desert]]s but have adapted to a wide range of environmental conditions, and can be found on all continents except [[Antarctica]]. There are over 2,500 described [[species]], with 22 extant (living) families recognized to date. Their [[Taxonomy (biology)|taxonomy]] is being revised to account for 21st-century [[genomic]] studies.
 Scorpions primarily prey on [[insect]]s and other [[invertebrate]]s, but some species hunt [[vertebrate]]s. They use their pincers to restrain and kill prey, or to prevent their own predation. The [[Scorpion sting|venomous sting]] is used for offense and defense. During courtship, the male and female grasp each other's pincers and dance while he tries to move her onto his [[sperm packet]]. All known species give [[viviparous|live birth]] and the female cares for the young as their [[exoskeleton]]s harden, transporting them on her back. The exoskeleton contains [[fluorescent]] chemicals and glows under [[ultraviolet]] light.
@@ Line 20: / Line 23: @@
 The vast majority of species do not seriously threaten humans, and healthy adults usually do not need medical treatment after a sting. About 25 species (fewer than one [[percent]]) have venom capable of killing a human, which happens frequently in the parts of the world where they live, primarily where access to medical treatment is unlikely.
-Scorpions appear in art, folklore, mythology, and commercial brands. [[Scorpion motif]]s are woven into [[kilim]] carpets for protection from their sting. [[Scorpius (constellation)|Scorpius]] is the name of a constellation; the corresponding [[Scorpio (astrology)|astrological sign is Scorpio]]. A classical myth about Scorpius tells how the giant scorpion and its enemy [[Orion (mythology)|Orion]] became constellations on opposite sides of the sky.
+Scorpions appear in art, folklore, mythology, and commercial brands. [[Scorpion motif]]s are woven into [[kilim]] carpets for protection from their sting. [[Scorpius]] is the name of a constellation; the corresponding [[Scorpio (astrology)|astrological sign is Scorpio]]. A classical myth about Scorpius tells how the giant scorpion and its enemy [[Orion (mythology)|Orion]] became constellations on opposite sides of the sky.
 ==Etymology==
-The word "[[:wikt:scorpion|scorpion]]" originated in [[Middle English]] between 1175 and 1225 AD from [[Old French]] ''{{lang|fro|scorpion}}'',<ref>{{cite web |title=Scorpion |website=[[The American Heritage Dictionary of the English Language]], 5th Edition |date=2016 |access-date=7 December 2020 |url=http://www.thefreedictionary.com/scorpion |archive-date=22 November 2020 |archive-url=https://web.archive.org/web/20201122213551/https://www.thefreedictionary.com/scorpion |url-status=live }}</ref> or from Italian ''{{Lang|it|scorpione}}'', both derived from the Latin ''{{Lang|la|scorpio}}'', equivalent to ''{{Lang|la|scorpius}}'',<ref>{{cite web |title=Scorpion |website=Dictionary.com |url=http://dictionary.reference.com/browse/scorpion |access-date=7 December 2020 |archive-date=15 November 2015 |archive-url=https://web.archive.org/web/20151115230956/http://dictionary.reference.com/browse/scorpion |url-status=live }}</ref> which is the [[romanization]] of the Greek {{lang|el|[[:wikt:σκορπίος|σκορπίος]]}}&nbsp;– ''{{lang|el|skorpíos}}'',<ref>{{LSJ|skorpi/os|σκορπίος|ref}}.</ref> ultimately from [[Proto-Indo-European root]] ''*(s)ker-'' meaning "to cut", cf. "[[wikt:shear|shear]]".<ref>{{OEtymD|scorpion}}</ref>
+The word ''scorpion'' originated in [[Middle English]] between 1175 and 1225 AD from [[Old French]] ''{{lang|fro|scorpion}}'',<ref>{{cite web |title=Scorpion |website=[[The American Heritage Dictionary of the English Language]], 5th Edition |date=2016 |access-date=7 December 2020 |url=http://www.thefreedictionary.com/scorpion |archive-date=22 November 2020 |archive-url=https://web.archive.org/web/20201122213551/https://www.thefreedictionary.com/scorpion |url-status=live }}</ref> or from Italian ''{{Lang|it|scorpione}}'', both derived from the Latin ''{{Lang|la|scorpio}}'', equivalent to ''{{Lang|la|scorpius}}'',<ref>{{cite web |title=Scorpion |website=Dictionary.com |url=http://dictionary.reference.com/browse/scorpion |access-date=7 December 2020 |archive-date=15 November 2015 |archive-url=https://web.archive.org/web/20151115230956/http://dictionary.reference.com/browse/scorpion |url-status=live }}</ref> which is the [[romanization]] of the Greek {{lang|el|[[:wikt:σκορπίος|σκορπίος]]}}&nbsp;– ''{{lang|el|skorpíos}}'',<ref>{{LSJ|skorpi/os|σκορπίος|ref}}.</ref> with no native IE etymology (cfr.  Arabic ''[[:wikt:العقرب|ʕaqrab]]'' 'scorpion', Proto-Germanic ''[[:wikt:crab#Etymology_1|*krabbô]]'' 'crab').
 ==Evolution==
@@ Line 31: / Line 34: @@
 [[File: PSM V27 D413 Fossil scorpion.jpg|thumb|''[[Palaeophonus|Palaeophonus nuncius]]'', a [[Silurian]] fossil from Sweden]]
-Scorpion [[fossil]]s have been found in many [[strata]], including marine [[Silurian]] and estuarine [[Devonian]] deposits, coal deposits from the [[Carboniferous Period]] and in [[amber]]. Whether the early scorpions were marine or terrestrial has been debated, though they had [[book lung]]s like modern terrestrial species.<ref name="Howard Edgecombe 2019"/><ref>{{cite journal |last1=Scholtz |first1=Gerhard |last2=Kamenz |first2=Carsten |title=The Book Lungs of Scorpiones and Tetrapulmonata (Chelicerata, Arachnida): Evidence for Homology and a Single Terrestrialisation Event of a Common Arachnid Ancestor |year=2006 |journal=Zoology |volume=109 |issue=1 |pages=2–13 |pmid=16386884 |doi=10.1016/j.zool.2005.06.003 }}</ref><ref>{{cite journal |last1=Dunlop |first1=Jason A. |last2=Tetlie |first2=O. Erik |last3=Prendini |first3=Lorenzo |doi=10.1111/j.1475-4983.2007.00749.x |title=Reinterpretation of the Silurian Scorpion ''Proscorpius osborni'' (Whitfield): Integrating Data from Palaeozoic and Recent Scorpions |year=2008 |journal=Palaeontology |volume=51 |issue=2 |pages=303–320|doi-access=free }}</ref><ref>{{cite journal |last1=Kühl |first1=G. |first2=A. |last2=Bergmann |first3=J. |last3=Dunlop |first4=R. J. |last4=Garwood |first5=J. |last5=Rust |doi=10.1111/j.1475-4983.2012.01152.x |title=Redescription and Palaeobiology of ''Palaeoscorpius devonicus'' Lehmann, 1944 from the Lower Devonian Hunsrück Slate of Germany |year=2012 |journal=Palaeontology |volume=55 |issue=4 |pages=775–787|doi-access=free }}</ref> Over 100 fossil species of scorpion have been described.<ref name=Siri>{{cite book |last1=Dunlop |first1=J. A. |last2=Penney |first2=D. |year=2012 |title=Fossil Arachnids |publisher=Siri Scientific Press |page=23 |isbn=978-0956779540}}</ref> The oldest found as of 2021 is ''[[Dolichophonus|Dolichophonus loudonensis]]'', which lived during the Silurian, in present-day Scotland.<ref>{{Cite journal|last1=Anderson|first1=Evan P.|last2=Schiffbauer|first2=James D.|last3=Jacquet|first3=Sarah M.|last4=Lamsdell|first4=James C.|last5=Kluessendorf|first5=Joanne|last6=Mikulic|first6=Donald G.|date=2021|title=Stranger than a scorpion: a reassessment of Parioscorpio venator, a problematic arthropod from the Llandoverian Waukesha Lagerstätte|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/pala.12534|journal=Palaeontology|language=en|volume=64|issue=3|pages=429–474|doi=10.1111/pala.12534|s2cid=234812878|issn=1475-4983|access-date=20 April 2021|archive-date=20 April 2021|archive-url=https://web.archive.org/web/20210420145034/https://onlinelibrary.wiley.com/doi/abs/10.1111/pala.12534|url-status=live}}</ref> ''[[Gondwanascorpio]]'' from the Devonian is among the earliest-known terrestrial animals on the [[Gondwana]] supercontinent.<ref>{{cite journal |last=Gess |first=R. W. |year=2013 |title=The Earliest Record of Terrestrial Animals in Gondwana: a Scorpion from the Famennian (Late Devonian) Witpoort Formation of South Africa |journal=[[African Invertebrates]] |volume=54 |issue=2 |pages=373–379 |url=http://africaninvertebrates.org/ojs/index.php/AI/article/view/284 |doi=10.5733/afin.054.0206 |doi-access=free |access-date=28 August 2013 |archive-date=6 September 2013 |archive-url=https://web.archive.org/web/20130906073206/http://africaninvertebrates.org/ojs/index.php/AI/article/view/284 |url-status=live }}</ref> Some Palaeozoic scorpions possessed [[compound eye]]s similar to those of eurypterids.<ref>[https://pubmed.ncbi.nlm.nih.gov/31780700/ Insights into the 400 million-year-old eyes of giant sea scorpions (Eurypterida) suggest the structure of Palaeozoic compound eyes]</ref> The [[Triassic]] fossils ''[[Protochactas]]'' and ''[[Protobuthus]]'' belong to the modern clades [[Chactoidea]] and [[Buthoidea]] respectively, indicating that the [[crown group]] of modern scorpions had emerged by this time.<ref>{{Cite journal |last1=Magnani, Fabio |last2=Stockar, Rudolf |last3=Lourenço, Wilson R. |date=2022 |others=Lionel Delaunay |title=Une nouvelle famille, genre et espèce de scorpion fossile du Calcaire de Meride (Trias Moyen) du Mont San Giorgio (Suisse)A new family, genus and species of fossil scorpion from the Meride Limestone (Middle Triassic) of Monte San Giorgio (Switzerland) |journal=Faunitaxys |volume=10 |issue=24 |url=https://hal.archives-ouvertes.fr/hal-03660445v1 |language=en |pages=1–7 |doi=10.57800/FAUNITAXYS-10(24)}}</ref>
+Scorpion [[fossil]]s have been found in many [[strata]], including marine [[Silurian]] and estuarine [[Devonian]] deposits, coal deposits from the [[Carboniferous Period]] and in [[amber]]. Whether the early scorpions were marine or terrestrial has been debated, though they had [[book lung]]s like modern terrestrial species.<ref name="Howard Edgecombe 2019"/><ref>{{cite journal |last1=Scholtz |first1=Gerhard |last2=Kamenz |first2=Carsten |title=The Book Lungs of Scorpiones and Tetrapulmonata (Chelicerata, Arachnida): Evidence for Homology and a Single Terrestrialisation Event of a Common Arachnid Ancestor |year=2006 |journal=Zoology |volume=109 |issue=1 |pages=2–13 |pmid=16386884 |doi=10.1016/j.zool.2005.06.003 |bibcode=2006Zool..109....2S }}</ref><ref>{{cite journal |last1=Dunlop |first1=Jason A. |last2=Tetlie |first2=O. Erik |last3=Prendini |first3=Lorenzo |doi=10.1111/j.1475-4983.2007.00749.x |title=Reinterpretation of the Silurian Scorpion ''Proscorpius osborni'' (Whitfield): Integrating Data from Palaeozoic and Recent Scorpions |year=2008 |journal=Palaeontology |volume=51 |issue=2 |pages=303–320|bibcode=2008Palgy..51..303D |doi-access=free }}</ref><ref>{{cite journal |last1=Kühl |first1=G. |first2=A. |last2=Bergmann |first3=J. |last3=Dunlop |first4=R. J. |last4=Garwood |first5=J. |last5=Rust |doi=10.1111/j.1475-4983.2012.01152.x |title=Redescription and Palaeobiology of ''Palaeoscorpius devonicus'' Lehmann, 1944 from the Lower Devonian Hunsrück Slate of Germany |year=2012 |journal=Palaeontology |volume=55 |issue=4 |pages=775–787|bibcode=2012Palgy..55..775K |doi-access=free }}</ref> Over 100 fossil species of scorpion have been described.<ref name=Siri>{{cite book |last1=Dunlop |first1=J. A. |last2=Penney |first2=D. |year=2012 |title=Fossil Arachnids |publisher=Siri Scientific Press |page=23 |isbn=978-0956779540}}</ref> The oldest found as of 2021 is ''[[Dolichophonus|Dolichophonus loudonensis]]'', which lived during the Silurian, in present-day Scotland.<ref>{{Cite journal|last1=Anderson|first1=Evan P.|last2=Schiffbauer|first2=James D.|last3=Jacquet|first3=Sarah M.|last4=Lamsdell|first4=James C.|last5=Kluessendorf|first5=Joanne|last6=Mikulic|first6=Donald G.|date=2021|title=Stranger than a scorpion: a reassessment of Parioscorpio venator, a problematic arthropod from the Llandoverian Waukesha Lagerstätte|url=https://onlinelibrary.wiley.com/doi/abs/10.1111/pala.12534|journal=Palaeontology|language=en|volume=64|issue=3|pages=429–474|doi=10.1111/pala.12534|bibcode=2021Palgy..64..429A |s2cid=234812878|issn=1475-4983|access-date=20 April 2021|archive-date=20 April 2021|archive-url=https://web.archive.org/web/20210420145034/https://onlinelibrary.wiley.com/doi/abs/10.1111/pala.12534|url-status=live}}</ref> ''[[Gondwanascorpio]]'' from the Devonian is among the earliest-known terrestrial animals on the [[Gondwana]] supercontinent.<ref>{{cite journal |last=Gess |first=R. W. |year=2013 |title=The Earliest Record of Terrestrial Animals in Gondwana: a Scorpion from the Famennian (Late Devonian) Witpoort Formation of South Africa |journal=[[African Invertebrates]] |volume=54 |issue=2 |pages=373–379 |url=http://africaninvertebrates.org/ojs/index.php/AI/article/view/284 |doi=10.5733/afin.054.0206 |doi-access=free |bibcode=2013AfrIn..54..373G |access-date=28 August 2013 |archive-date=6 September 2013 |archive-url=https://web.archive.org/web/20130906073206/http://africaninvertebrates.org/ojs/index.php/AI/article/view/284 |url-status=live }}</ref> Some Palaeozoic scorpions possessed [[compound eye]]s similar to those of eurypterids.<ref>{{cite journal | pmid=31780700 | year=2019 | last1=Schoenemann | first1=B. | last2=Poschmann | first2=M. | author3=Clarkson ENK | title=Insights into the 400 million-year-old eyes of giant sea scorpions (Eurypterida) suggest the structure of Palaeozoic compound eyes | journal=Scientific Reports | volume=9 | issue=1 | page=17797 | doi=10.1038/s41598-019-53590-8 | pmc=6882788 | bibcode=2019NatSR...917797S }}</ref> The [[Triassic]] fossils ''[[Protochactas]]'' and ''[[Protobuthus]]'' belong to the modern clades [[Chactoidea]] and [[Buthoidea]] respectively, indicating that the [[crown group]] of modern scorpions had emerged by this time.<ref>{{Cite journal |last1=Magnani, Fabio |last2=Stockar, Rudolf |last3=Lourenço, Wilson R. |date=2022 |others=Lionel Delaunay |title=Une nouvelle famille, genre et espèce de scorpion fossile du Calcaire de Meride (Trias Moyen) du Mont San Giorgio (Suisse)A new family, genus and species of fossil scorpion from the Meride Limestone (Middle Triassic) of Monte San Giorgio (Switzerland) |journal=Faunitaxys |volume=10 |issue=24 |url=https://hal.archives-ouvertes.fr/hal-03660445v1 |language=en |pages=1–7 |doi=10.57800/FAUNITAXYS-10(24)}}</ref>
 ===Phylogeny===
@@ Line 39: / Line 42: @@
 |label1=[[Chelicerata]]
 |1={{clade
-   |1=[[Pycnogonida]] (sea spiders) [[File:Pycnogonida Nymphon s Sars (white background).png|80px]]
+   |1=[[Pycnogonida]] (sea spiders) [[File:Nymphon signatum 194389384 (white background).jpg|90px]]
    |label2=[[Prosomapoda]]
    |2={{clade
-      |1=[[Xiphosura]] (horseshoe crabs) [[File:FMIB 51225 Horse-Shoe Crab, Limulus Polyphemus, Latreille (cropped).jpeg|50px]]
+      |1=[[Xiphosura]] (horseshoe crabs) [[File:Limulus polyphemus (aquarium) (white background).jpg|70px]]
       |2={{clade
-         |1=†[[Eurypterida]] (sea scorpions) [[File:Hibbertopterus scouleri.jpg|80px]]
+         |1=†[[Eurypterida]] (sea scorpions) <span style="{{MirrorH}}">[[File:Eurypterus Paleoart (no background).png|70 px]]</span>
          |label2=[[Arachnida]]
          |2={{clade
             |label1=Non-pulmonates
-            |1=([[ticks]], [[harvestmen]], etc) [[File:Trombidium holosericeum (white background).jpg|40px]]
+            |1=([[ticks]], [[harvestmen]], etc) [[File:Ixodes scapularis P1170301a (white background).png|70px]]
             |label2=pulmonates
             |2={{clade
                |label1='''Scorpiones'''
-               |1=[[File:Chaerilus pseudoconchiformus male (cropped).jpg|50px]]
+               |1=[[File:Buthus_mariefranceae_(10.3897-zookeys.686.12206)_Figure_1.jpg|65px]]
                |label2=[[Tetrapulmonata]]
                |2={{clade
-                  |1=[[Araneae]] (spiders) [[File:Theraphosa blondi MHNT.jpg|50px]]
+                  |1=[[Araneae]] (spiders) [[File:Aptostichus simus Monterey County.jpg|70px]]
-                  |2=Pedipalpi ([[whip scorpions]], etc) [[File:Damon johnstoni – Lydekker, 1879.png|80px]]
+                  |2=Pedipalpi ([[whip scorpions]], etc) [[File:Whip Scorpion body (9672115742) (white background).png|80px]]
                   }}
                }}
@@ Line 65: / Line 68: @@
 }}
-Recent studies place [[pseudoscorpion]]s as the sister group of scorpions in the clade Panscorpiones, which together with Tetrapulmonata makes up the clade Arachnopulmonata.<ref>[https://www.biorxiv.org/content/10.1101/2020.11.18.389098v1.full Taxonomic sampling and rare genomic changes overcome long-branch attraction in the phylogenetic placement of pseudoscorpions]</ref>
+Recent studies place [[pseudoscorpion]]s as the sister group of scorpions in the clade Panscorpiones, which together with Tetrapulmonata makes up the clade Arachnopulmonata.<ref>{{Cite journal|url=https://www.biorxiv.org/content/10.1101/2020.11.18.389098v1|title=Taxonomic sampling and rare genomic changes overcome long-branch attraction in the phylogenetic placement of pseudoscorpions|first1=Andrew Z.|last1=Ontano|first2=Guilherme|last2=Gainett|first3=Shlomi|last3=Aharon|first4=Jesús A.|last4=Ballesteros|first5=Ligia R.|last5=Benavides|first6=Kevin F.|last6=Corbett|first7=Efrat|last7=Gavish-Regev|first8=Mark S.|last8=Harvey|first9=Scott|last9=Monsma|first10=Carlos E.|last10=Santibáñez-López|first11=Emily V. W.|last11=Setton|first12=Jakob T.|last12=Zehms|first13=Jeanne A.|last13=Zeh|first14=David W.|last14=Zeh|first15=Prashant P.|last15=Sharma|journal=Molecular Biology and Evolution|volume=38|issue=6|date=June 2021|pages=2446–2467|doi=10.1093/molbev/msab038|pmid=33565584 |pmc=8136511 }}</ref>
-The internal [[phylogeny]] of the scorpions has been debated,<ref name="Howard Edgecombe 2019">{{cite journal |last1=Howard |first1=Richard J. |last2=Edgecombe |first2=Gregory D. |last3=Legg |first3=David A. |last4=Pisani |first4=Davide |last5=Lozano-Fernandez |first5=Jesus |title=Exploring the Evolution and Terrestrialization of Scorpions (Arachnida: Scorpiones) with Rocks and Clocks |journal=Organisms Diversity & Evolution |volume=19 |issue=1 |year=2019 |pages=71–86 |issn=1439-6092 |doi=10.1007/s13127-019-00390-7 |doi-access=free}}</ref> but [[genomic]] analysis consistently places the [[Bothriuridae]] as sister to a clade consisting of Scorpionoidea and [[Chactoidea]]. The scorpions diversified between the Devonian and the early [[Carboniferous]]. The main division is into the clades Buthida and Iurida. The Bothriuridae diverged starting before temperate Gondwana broke up into separate land masses, completed by the [[Jurassic]]. The Iuroidea and Chactoidea are both seen not to be single clades, and are shown as "[[paraphyletic]]" (with quotation marks) in this 2018 cladogram.<ref name="Sharma Baker 2018">{{cite journal |last1=Sharma |first1=Prashant P. |last2=Baker |first2=Caitlin M. |last3=Cosgrove |first3=Julia G. |last4=Johnson |first4=Joanne E. |last5=Oberski |first5=Jill T. |last6=Raven |first6=Robert J. |last7=Harvey |first7=Mark S. |last8=Boyer |first8=Sarah L. |last9=Giribet |first9=Gonzalo |title=A Revised Dated Phylogeny of Scorpions: Phylogenomic Support for Ancient Divergence of the Temperate Gondwanan Family Bothriuridae |journal=Molecular Phylogenetics and Evolution |volume=122 |year=2018 |pages=37–45 |issn=1055-7903 |doi=10.1016/j.ympev.2018.01.003 |pmid=29366829 }}</ref>
+The internal [[phylogeny]] of the scorpions has been debated,<ref name="Howard Edgecombe 2019">{{cite journal |last1=Howard |first1=Richard J. |last2=Edgecombe |first2=Gregory D. |last3=Legg |first3=David A. |last4=Pisani |first4=Davide |last5=Lozano-Fernandez |first5=Jesus |title=Exploring the Evolution and Terrestrialization of Scorpions (Arachnida: Scorpiones) with Rocks and Clocks |journal=Organisms Diversity & Evolution |volume=19 |issue=1 |year=2019 |pages=71–86 |issn=1439-6092 |doi=10.1007/s13127-019-00390-7 |doi-access=free|hdl=10261/217081 |hdl-access=free }}</ref> but [[genomic]] analysis consistently places the [[Bothriuridae]] as sister to a clade consisting of Scorpionoidea and [[Chactoidea]]. The scorpions diversified between the Devonian and the early [[Carboniferous]]. The main division is into the clades Buthida and Iurida. The Bothriuridae diverged starting before temperate Gondwana broke up into separate land masses, completed by the [[Jurassic]]. The Iuroidea and Chactoidea are both seen not to be single clades, and are shown as "[[paraphyletic]]" (with quotation marks) in this 2018 cladogram.<ref name="Sharma Baker 2018">{{cite journal |last1=Sharma |first1=Prashant P. |last2=Baker |first2=Caitlin M. |last3=Cosgrove |first3=Julia G. |last4=Johnson |first4=Joanne E. |last5=Oberski |first5=Jill T. |last6=Raven |first6=Robert J. |last7=Harvey |first7=Mark S. |last8=Boyer |first8=Sarah L. |last9=Giribet |first9=Gonzalo |title=A Revised Dated Phylogeny of Scorpions: Phylogenomic Support for Ancient Divergence of the Temperate Gondwanan Family Bothriuridae |journal=Molecular Phylogenetics and Evolution |volume=122 |year=2018 |pages=37–45 |issn=1055-7903 |doi=10.1016/j.ympev.2018.01.003 |pmid=29366829 |doi-access=free |bibcode=2018MolPE.122...37S }}</ref>
 {{clade
@@ Line 78: / Line 81: @@
          |2=[[Pseudochactoidea]] <!--(9 spp.)--> [[File:Vietbocap canhi (white background).jpg|45px]]
          }}
-      |2=[[Buthoidea]] <!--(1209 spp.)--> [[File:1412849934052m04SlwWVfJyEgvBO Buthidae (white background).jpg|50px]]
+      |2=[[Buthoidea]] <!--(1209 spp.)--> [[File:Buthus mariefranceae (10.3897-zookeys.686.12206) Figure 1.jpg|65px]]
       }}
    |label2=Iurida<!--Parvorder-->
@@ Line 93: / Line 96: @@
                |2="[[Chactoidea]]" (part)
                }}
-            |2=[[Scorpionoidea]] <!--(183 spp.)--> [[File:Female Emperor Scorpion.jpg|75px]]
+            |2=[[Scorpionoidea]] <!--(183 spp.)--> <span style="{{MirrorH}}">[[File:Female Emperor Scorpion.jpg|75px]]</span>
             }}
          }}
@@ Line 103: / Line 106: @@
 {{main|Taxonomy of scorpions}}
-[[Carl Linnaeus]] described six species of scorpion in his genus ''Scorpio'' in 1758 and 1767; three of these are now considered valid and are called ''[[Scorpio maurus]]'', ''[[Androctonus australis]]'', and ''[[Euscorpius]] carpathicus''; the other three are dubious names. He placed the scorpions among his "Insecta aptera" (wingless insects), a group that included Crustacea, Arachnida and [[Myriapoda]].<ref>{{cite journal |last1=Fet |first1=V. |last2=Braunwalder |first2=M. E. |last3=Cameron |first3=H. D. |title=Scorpions (Arachnida, Scorpiones) Described by Linnaeus |journal=Bulletin of the British Arachnological Society |date=2002 |volume=12 |issue=4 |pages=176–182 |url=http://britishspiders.org.uk/bulletin/120405.pdf |access-date=20 October 2020 |archive-date=26 September 2020 |archive-url=https://web.archive.org/web/20200926182646/http://britishspiders.org.uk/bulletin/120405.pdf |url-status=live }}</ref> In 1801, [[Jean-Baptiste Lamarck]] divided up the "Insecta aptera", creating the [[taxon]] Arachnides for spiders, scorpions, and acari (mites and ticks), though it also contained the [[Thysanura]] (thrips), Myriapoda and parasites such as lice.<ref name="Burmeister 1836">{{cite book |last1=Burmeister |first1=Carl Hermann C. |last2=Shuckard |first2=W. E. (trans) |title=A Manual of Entomology |url=https://books.google.com/books?id=g9sDAAAAQAAJ&pg=PA613 |year=1836 |pages=613ff |access-date=20 October 2020 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031222/https://books.google.com/books?id=g9sDAAAAQAAJ&pg=PA613 |url-status=live }}</ref> German [[arachnologist]] [[Carl Ludwig Koch]] created the order Scorpiones in 1837. He divided it into four families, the six-eyed scorpions "Scorpionides", the eight-eyed scorpions "Buthides", the ten-eyed scorpions "Centrurides", and the twelve-eyed scorpions "Androctonides".<ref>{{cite book |last=Koch |first=Carl Ludwig |title=Übersicht des Arachnidensystems |date=1837 |publisher=C. H. Zeh |pages=86–92 |url=https://archive.org/details/bersichtdesara1518371850koch/page/n91/mode/2up?q=Scorpiones |language=de}}</ref>
+[[Carl Linnaeus]] described six species of scorpion in his genus ''Scorpio'' in 1758 and 1767; three of these are now considered valid and are called ''[[Scorpio maurus]]'', ''[[Androctonus australis]]'', and ''[[Euscorpius]] carpathicus''; the other three are dubious names. He placed the scorpions among his "Insecta aptera" (wingless insects), a group that included Crustacea, Arachnida and [[Myriapoda]].<ref>{{cite journal |last1=Fet |first1=V. |last2=Braunwalder |first2=M. E. |last3=Cameron |first3=H. D. |title=Scorpions (Arachnida, Scorpiones) Described by Linnaeus |journal=Bulletin of the British Arachnological Society |date=2002 |volume=12 |issue=4 |pages=176–182 |url=http://britishspiders.org.uk/bulletin/120405.pdf |access-date=20 October 2020 |archive-date=26 September 2020 |archive-url=https://web.archive.org/web/20200926182646/http://britishspiders.org.uk/bulletin/120405.pdf |url-status=live }}</ref> In 1801, [[Jean-Baptiste Lamarck]] divided up the "Insecta aptera", creating the [[taxon]] Arachnides for spiders, scorpions, and acari (mites and ticks), though it also contained the [[Thysanura]], Myriapoda and parasites such as lice.<ref name="Burmeister 1836">{{cite book |last1=Burmeister |first1=Carl Hermann C. |last2=Shuckard |first2=W. E. (trans) |title=A Manual of Entomology |url=https://books.google.com/books?id=g9sDAAAAQAAJ&pg=PA613 |year=1836 |pages=613ff |access-date=20 October 2020 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031222/https://books.google.com/books?id=g9sDAAAAQAAJ&pg=PA613 |url-status=live }}</ref> German [[arachnologist]] [[Carl Ludwig Koch]] created the order Scorpiones in 1837. He divided it into four families, the six-eyed scorpions "Scorpionides", the eight-eyed scorpions "Buthides", the ten-eyed scorpions "Centrurides", and the twelve-eyed scorpions "Androctonides".<ref>{{cite book |last=Koch |first=Carl Ludwig |title=Übersicht des Arachnidensystems |date=1837 |publisher=C. H. Zeh |pages=86–92 |url=https://archive.org/details/bersichtdesara1518371850koch/page/n91/mode/2up?q=Scorpiones |language=de}}</ref>
 More recently, some twenty-two families containing over 2,500 species of scorpions have been described, with many additions and much reorganization of taxa in the 21st century.<ref name="Rein 2020"/><ref name="Howard Edgecombe 2019"/><ref>{{cite web |url=http://kovarex.com/scorpio/pdf/scorpions-Kovarik-2009.pdf |first=František |last=Kovařík |year=2009 |title=Illustrated Catalog of Scorpions, Part I |access-date=22 January 2011 |archive-date=26 October 2010 |archive-url=https://web.archive.org/web/20101026163042/http://kovarex.com/scorpio/pdf/scorpions-Kovarik-2009.pdf |url-status=live }}</ref> There are over 100 described taxa of fossil scorpions.<ref name=Siri/> This classification is based on Soleglad and Fet (2003),<ref>{{cite journal |last1=Soleglad |first1=Michael E. |last2=Fet |first2=Victor |year=2003 |title=High-level Systematics and Phylogeny of the Extant Scorpions (Scorpiones: Orthosterni) |journal=[[Euscorpius (journal)|Euscorpius]] |volume=11 |pages=1–175 |url=http://www.science.marshall.edu/fet/euscorpius/pubs.htm |format=multiple parts |access-date=13 June 2008 |archive-date=8 December 2019 |archive-url=https://web.archive.org/web/20191208071547/http://www.science.marshall.edu/fet/euscorpius/pubs.htm |url-status=live }}</ref> which replaced Stockwell's older, unpublished classification.<ref>{{cite thesis |last=Stockwell |first=Scott A. |title=Revision of the Phylogeny and Higher Classification of Scorpions (Chelicerata) |year=1989 |publisher=[[University of California, Berkeley]]|type=PhD thesis}}</ref> Further taxonomic changes are from papers by Soleglad et al. (2005).<ref name=Soleglad2005>{{cite journal |last1=Soleglad |first1=Michael E. |last2=Fet |first2=Victor |last3=Kovařík |first3=F. |year=2005 |title=The Systematic Position of the Scorpion Genera ''Heteroscorpion'' Birula, 1903 and ''Urodacus'' Peters, 1861 (Scorpiones: Scorpionoidea) |journal=[[Euscorpius (journal)|Euscorpius]] |volume=20 |pages=1–38 |url=http://www.science.marshall.edu/fet/euscorpius/p2005_20.pdf |access-date=13 June 2008 |archive-date=16 December 2008 |archive-url=https://web.archive.org/web/20081216234048/http://www.science.marshall.edu/fet/euscorpius/p2005_20.pdf |url-status=live }}</ref><ref>{{cite journal |last1=Fet |first1=Victor |last2=Soleglad |first2=Michael E. |year=2005 |title=Contributions to Scorpion Systematics. I. On Recent Changes in High-level Taxonomy |journal=[[Euscorpius (journal)|Euscorpius]] |issue=31 |pages=1–13 |url=http://www.science.marshall.edu/fet/euscorpius/p2005_31.pdf |issn=1536-9307 |access-date=7 April 2010 |archive-date=26 April 2018 |archive-url=https://web.archive.org/web/20180426135453/http://www.science.marshall.edu/fet/euscorpius/p2005_31.pdf |url-status=live }}</ref>
@@ Line 121: / Line 124: @@
 * Parvorder [[Buthida]] <small>[[Michael E. Soleglad|Soleglad]] & [[Victor Fet|Fet]], 2003</small>
 <!----------------------------------------------->
-** Superfamily [[Buthoidea]] <small>[[Carl Ludwig Koch|C. L. Koch]], 1837</small>
+** Superfamily [[Buthoidea]] <small>[[C. L. Koch]], 1837</small>
-*** Family [[Buthidae]] <small>[[Carl Ludwig Koch|C. L. Koch]], 1837</small> (1209 spp.) (thick-tailed scorpions, including the most dangerous species)
+*** Family [[Buthidae]] <small>[[C. L. Koch]], 1837</small> (1209 spp.) (thick-tailed scorpions, including the most dangerous species)
 *** Family [[Microcharmidae]] <small>[[Wilson R. Lourenço|Lourenço]], 1996, 2019</small> (17 spp.) (African scorpions of humid forest leaf litter)
 <!----------------------------------------------->
@@ Line 148: / Line 151: @@
 *** Family [[Iuridae]] <small>[[Tord Tamerlan Teodor Thorell|Thorell]], 1876</small> (21 spp.) (scorpions with a large tooth on inner side of moveable claw)
 <!----------------------------------------------->
-** Superfamily [[Scorpionoidea]] <small>[[Pierre André Latreille|Latreille]], 1802</small>
+** Superfamily [[Scorpionoidea]] <small>[[Latreille]], 1802</small>
 *** Family [[Bothriuridae]] <small>[[Eugène Simon|Simon]], 1880</small> (158 spp.) (Southern hemisphere tropical and temperate scorpions)
 *** Family [[Hemiscorpiidae]] <small>[[Reginald Innes Pocock|Pocock]], 1893</small> (16 spp.) (rock, creeping, or tree scorpions of the Middle East)
 *** Family [[Hormuridae]] <small>[[Malcolm Laurie|Laurie]], 1896</small> (92 spp.) (flattened, crevice-living scorpions of Southeast Asia and Australia)
 *** Family [[Rugodentidae]] <small>Bastawade et al., 2005</small> (1 sp.) (burrowing scorpions of India)
-*** Family [[Scorpionidae]] <small>[[Pierre André Latreille|Latreille]], 1802</small> (183 spp.) (burrowing or pale-legged scorpions)
+*** Family [[Scorpionidae]] <small>[[Latreille]], 1802</small> (183 spp.) (burrowing or pale-legged scorpions)
 *** Family [[Diplocentridae]] <small>[[Ferdinand Karsch|Karsch]], 1880</small> (134 spp.) (closely related to and sometimes placed in Scorpionidae, but have spine on telson)
 *** Family [[Heteroscorpionidae]] <small>[[Karl Kraepelin|Kraepelin]], 1905</small> (6 spp.) (scorpions of Madagascar)
@@ Line 160: / Line 163: @@
 == Geographical distribution ==
-Scorpions are found on all continents except [[Antarctica]]. The diversity of scorpions is greatest in subtropical areas; it decreases toward the poles and equator, though scorpions are found in the tropics. Scorpions did not occur naturally in [[Great Britain]] but were accidentally [[Introduced species|introduced]] by humans, and have now established a population.{{sfn|Polis|1990|p=249}}<ref>{{Cite web |last=Nurden |first=John |date=2021-07-05 |title=We track down the UK's biggest scorpion colony |url=https://www.kentonline.co.uk/sheerness/news/we-track-down-the-uks-biggest-scorpion-colony-250024/ |access-date=2022-09-05 |website=Kent Online |language=en}}</ref> [[New Zealand]], and some of the islands in [[Oceania]], have in the past had small populations of introduced scorpions, but they were exterminated.<ref>{{Cite web |date=2018-04-06 |title=Critter of the Week The Pseudoscorpion! |url=https://www.rnz.co.nz/national/programmes/afternoons/audio/2018639383/critter-of-the-week-the-pseudoscorpion |access-date=2022-09-05 |website=RNZ |language=en-nz}}</ref>{{sfn|Polis|1990|p=249}} Five colonies of ''[[Euscorpius flavicaudis]]'' have established themselves since the late 19th century in [[Sheerness]] in England at 51°N,<ref>{{cite journal |first=T. G. |last=Benton |year=1992 |title=The Ecology of the Scorpion ''Euscorpius flavicaudis'' in England |journal=[[Journal of Zoology]] |volume=226 |issue=3 |pages=351–368 |doi=10.1111/j.1469-7998.1992.tb07484.x}}</ref><ref>{{cite journal |last1=Benton |first1=T. G. |title=The Life History of ''Euscorpius flavicaudis'' (Scorpiones, Chactidae) |journal=The Journal of Arachnology |date=1991 |volume=19 |issue=2 |pages=105–110 |jstor=3705658 |url=http://www.americanarachnology.org/JoA_free/JoA_v19_n2%20/JoA_v19_p105.pdf |access-date=16 January 2008 |archive-date=28 February 2008 |archive-url=https://web.archive.org/web/20080228194248/http://www.americanarachnology.org/JoA_free/JoA_v19_n2%20/JoA_v19_p105.pdf |url-status=live }}</ref><ref>{{cite web |url=https://www.ntnu.no/ub/scorpion-files/e_flavicaudis.php |title=''Euscorpius flavicaudis'' |access-date=13 June 2008 |first=Jan Ove |last=Rein |year=2000 |website=The Scorpion Files |publisher=[[Norwegian University of Science and Technology]] |archive-date=26 April 2018 |archive-url=https://web.archive.org/web/20180426135809/https://www.ntnu.no/ub/scorpion-files/e_flavicaudis.php |url-status=live }}</ref> while ''[[Paruroctonus boreus]]'' lives as far north as [[Red Deer, Alberta]], at 52°N.<ref>{{cite book |last=Johnson |first=D. L. |url=http://people.uleth.ca/~dan.johnson/scorpions/northern_scorpion_dj.pdf |date=2004 |chapter=The Northern Scorpion, ''Paruroctonus boreus'', in Southern Alberta, 1983–2003 |title=Arthropods of Canadian Grasslands 10 |publisher=Biological Survey of Canada |url-status=dead |archive-url=https://web.archive.org/web/20151129125229/http://people.uleth.ca/~dan.johnson/scorpions/northern_scorpion_dj.pdf |archive-date=29 November 2015}}</ref> A few species are on the [[IUCN Red List]]; ''[[Lychas braueri]]'' is classed as [[critically endangered]] (2014), ''[[Isometrus deharvengi]]'' as [[Endangered species|endangered]] (2016) and ''[[Chiromachus ochropus]]'' as [[Vulnerable species|vulnerable]] (2014).<ref>{{cite iucn |last=Gerlach |first=J. |year=2014 |title=''Lychas braueri''|url=https://www.iucnredlist.org/species/21460801/21567809 |access-date=10 December 2020}}</ref><ref>{{cite iucn |last1=Deharveng |first1=L. |last2=Bedos |first2=A. |year=2016 |title=''Isometrus deharvengi'' |url=https://www.iucnredlist.org/species/89656504/89656508 |access-date=10 December 2020}}</ref><ref>{{cite iucn |last=Gerlach |first=J. |year=2014 |title=''Chiromachus ochropus'' |url=https://www.iucnredlist.org/species/196784/21568706 |access-date=10 December 2020}}</ref>
+Scorpions are found on all continents except [[Antarctica]]. The diversity of scorpions is greatest in subtropical areas; it decreases toward the poles and equator, though scorpions are found in the tropics. Scorpions did not occur naturally in [[Great Britain]] but were accidentally [[Introduced species|introduced]] by humans, and have now established a population.{{sfn|Polis|1990|p=249}}<ref>{{Cite web |last=Nurden |first=John |date=2021-07-05 |title=We track down the UK's biggest scorpion colony |url=https://www.kentonline.co.uk/sheerness/news/we-track-down-the-uks-biggest-scorpion-colony-250024/ |access-date=2022-09-05 |website=Kent Online |language=en}}</ref> [[New Zealand]], and some of the islands in [[Oceania]], have in the past had small populations of introduced scorpions, but they were exterminated.<ref>{{Cite web |date=2018-04-06 |title=Critter of the Week The Pseudoscorpion! |url=https://www.rnz.co.nz/national/programmes/afternoons/audio/2018639383/critter-of-the-week-the-pseudoscorpion |access-date=2022-09-05 |website=RNZ |language=en-nz}}</ref>{{sfn|Polis|1990|p=249}} Five colonies of ''[[Euscorpius flavicaudis]]'' have established themselves since the late 19th century in [[Sheerness]] in England at 51°N,<ref>{{cite journal |first=T. G. |last=Benton |year=1992 |title=The Ecology of the Scorpion ''Euscorpius flavicaudis'' in England |journal=[[Journal of Zoology]] |volume=226 |issue=3 |pages=351–368 |doi=10.1111/j.1469-7998.1992.tb07484.x}}</ref><ref>{{cite journal |last1=Benton |first1=T. G. |title=The Life History of ''Euscorpius flavicaudis'' (Scorpiones, Chactidae) |journal=The Journal of Arachnology |date=1991 |volume=19 |issue=2 |pages=105–110 |jstor=3705658 |url=http://www.americanarachnology.org/JoA_free/JoA_v19_n2%20/JoA_v19_p105.pdf |access-date=16 January 2008 |archive-date=28 February 2008 |archive-url=https://web.archive.org/web/20080228194248/http://www.americanarachnology.org/JoA_free/JoA_v19_n2%20/JoA_v19_p105.pdf |url-status=live }}</ref><ref>{{cite web |url=https://www.ntnu.no/ub/scorpion-files/e_flavicaudis.php |title=''Euscorpius flavicaudis'' |access-date=13 June 2008 |first=Jan Ove |last=Rein |year=2000 |website=The Scorpion Files |publisher=[[Norwegian University of Science and Technology]] |archive-date=26 April 2018 |archive-url=https://web.archive.org/web/20180426135809/https://www.ntnu.no/ub/scorpion-files/e_flavicaudis.php |url-status=live }}</ref> while ''[[Paruroctonus boreus]]'' lives as far north as [[Red Deer, Alberta]], at 52°N.<ref>{{cite book |last=Johnson |first=D. L. |url=http://people.uleth.ca/~dan.johnson/scorpions/northern_scorpion_dj.pdf |date=2004 |chapter=The Northern Scorpion, ''Paruroctonus boreus'', in Southern Alberta, 1983–2003 |title=Arthropods of Canadian Grasslands 10 |publisher=Biological Survey of Canada |url-status=dead |archive-url=https://web.archive.org/web/20151129125229/http://people.uleth.ca/~dan.johnson/scorpions/northern_scorpion_dj.pdf |archive-date=29 November 2015}}</ref> A few species are on the [[IUCN Red List]]; ''[[Lychas braueri]]'' is classed as [[critically endangered]] (2014), ''[[Isometrus deharvengi]]'' as [[endangered]] (2016) and ''[[Chiromachus ochropus]]'' as [[Vulnerable species|vulnerable]] (2014).<ref>{{cite iucn |last=Gerlach |first=J. |year=2014 |title=''Lychas braueri''|url=https://www.iucnredlist.org/species/21460801/21567809 |access-date=10 December 2020}}</ref><ref>{{cite iucn |last1=Deharveng |first1=L. |last2=Bedos |first2=A. |year=2016 |title=''Isometrus deharvengi'' |url=https://www.iucnredlist.org/species/89656504/89656508 |access-date=10 December 2020}}</ref><ref>{{cite iucn |last=Gerlach |first=J. |year=2014 |title=''Chiromachus ochropus'' |url=https://www.iucnredlist.org/species/196784/21568706 |access-date=10 December 2020}}</ref>
 Scorpions are [[xerocole]]s, meaning they primarily live in [[desert]]s, but they can be found in virtually every terrestrial [[habitat]] including high-elevation mountains, caves, and [[intertidal zone]]s. They are largely absent from [[boreal ecosystem]]s such as the [[tundra]], high-altitude [[taiga]], and mountain tops.{{sfn|Polis|1990|pp=251–253}}<ref name="Howard Edgecombe 2019"/> The highest altitude reached by a scorpion is {{convert|5500|m|ft|sp=us}} in the Andes, for ''[[Orobothriurus crassimanus]]''.{{sfn|Stockmann|Ythier|2010|p=151}}
@@ Line 172: / Line 175: @@
 ===Cephalothorax===
-The cephalothorax comprises the [[carapace]], eyes, chelicerae (mouth parts), [[pedipalp]]s (which have [[chela (organ)|chelae]], commonly called claws or pincers) and four pairs of [[walking leg]]s. Scorpions have two eyes on the top of the cephalothorax, and usually two to five pairs of eyes along the front corners of the cephalothorax. While unable to form sharp images, their central eyes are amongst the most light sensitive in the animal kingdom, especially in dim light, and makes it possible for nocturnal species to use starlight to navigate at night.<ref name="Chakravarthy Sridhara 2016">{{cite book |last1=Chakravarthy |first1=Akshay Kumar |last2=Sridhara |first2=Shakunthala |title=Arthropod Diversity and Conservation in the Tropics and Sub-tropics |url=https://books.google.com/books?id=wLiVDQAAQBAJ&pg=PA60 |year=2016 |publisher=Springer |isbn=978-981-10-1518-2 |page=60 |access-date=11 September 2017 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031224/https://books.google.com/books?id=wLiVDQAAQBAJ&pg=PA60 |url-status=live }}</ref> The chelicerae are at the front and underneath the carapace. They are pincer-like and have three segments and sharp "teeth".{{sfn|Polis|1990|pp=16–17}}{{sfn|Stockmann|2015|p=26}} The brain of a scorpion is in the back of the cephalothorax, just above the [[esophagus]].{{sfn|Polis|1990|p=38}} As in other arachnids, the nervous system is highly concentrated in the cephalothorax, but has a long ventral nerve cord with segmented [[ganglia]] which may be a [[Primitive (phylogenetics)|primitive]] trait.{{sfn|Polis|1990|p=342}}
+The cephalothorax comprises the [[carapace]], eyes, chelicerae (mouth parts), [[pedipalp]]s (which have [[chelae]], commonly called claws or pincers) and four pairs of [[walking leg]]s. Scorpions have two eyes on the top of the cephalothorax, and usually two to five pairs of eyes along the front corners of the cephalothorax. While unable to form sharp images, their central eyes are amongst the most light sensitive in the animal kingdom, especially in dim light, which makes it possible for nocturnal species to use starlight to navigate at night.<ref name="Chakravarthy Sridhara 2016">{{cite book |last1=Chakravarthy |first1=Akshay Kumar |last2=Sridhara |first2=Shakunthala |title=Arthropod Diversity and Conservation in the Tropics and Sub-tropics |url=https://books.google.com/books?id=wLiVDQAAQBAJ&pg=PA60 |year=2016 |publisher=Springer |isbn=978-981-10-1518-2 |page=60 |access-date=11 September 2017 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031224/https://books.google.com/books?id=wLiVDQAAQBAJ&pg=PA60 |url-status=live }}</ref> The chelicerae are at the front and underneath the carapace. They are pincer-like and have three segments and sharp "teeth".{{sfn|Polis|1990|pp=16–17}}{{sfn|Stockmann|2015|p=26}} The brain of a scorpion is in the back of the cephalothorax, just above the [[esophagus]].{{sfn|Polis|1990|p=38}} As in other arachnids, the nervous system is highly concentrated in the cephalothorax, but has a long ventral nerve cord with segmented [[ganglia]] which may be a [[Primitive (phylogenetics)|primitive]] trait.{{sfn|Polis|1990|p=342}}
 The pedipalp is a segmented, clawed [[appendage]] used for prey immobilization, defense and sensory purposes. The segments of the pedipalp (from closest to the body outward) are coxa, trochanter, femur, patella, tibia (including the fixed claw and the manus) and tarsus (moveable claw). A scorpion has darkened or granular raised linear ridges, called "keels" or "carinae" on the pedipalp segments and on other parts of the body; these are useful as taxonomic [[phenotypic trait|characters]].{{sfn|Polis|1990|p=12}} Unlike those of some other arachnids, the legs have not been modified for other purposes, though they may occasionally be used for digging, and females may use them to catch emerging young. The legs are covered in [[proprioceptor]]s, [[bristle]]s and sensory [[setae]].{{sfn|Polis|1990|p=20}} Depending on the species, the legs may have spines and spurs.{{sfn|Polis|1990|p=74}}
 ===Mesosoma===
-[[File:Vista ventral de un escorpión.jpg|thumb|upright|right<!--left-->|Ventral view: the [[Pecten (biology)|pectines]] have a comblike structure in an inverted V shape.]]
+[[File:Vista ventral de un escorpión.jpg|thumb|upright|right<!--left-->|Ventral view: the [[pectines]] have a comblike structure in an inverted V shape.]]
-The mesosoma or preabdomen is the broad part of the opisthosoma.{{sfn|Polis|1990|pp=10–11}} In the early stages of embryonic development the mesosoma consist of eight segments, but the first segment disappear before birth, so the mesosoma in scorpions actually consist of segments 2-8.<ref>[https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3199777/ The ultrastructure of book lung development in the bark scorpion Centruroides gracilis (Scorpiones: Buthidae) - NCBI]</ref><ref>[http://www.sharmalabuw.org/uploads/1/3/6/1/13619635/schwager_et_al._2015.pdf Chelicerata - Prashant P. Sharma]</ref><ref>[http://www.science.marshall.edu/fet/euscorpius/p2011_120.pdf Euscorpius - Marshall University]</ref> These [[anterior]] seven [[somite]]s (segments) of the opisthosoma are each covered [[Anatomical terms of location|dorsally]] by a sclerotized plate called the [[tergum|tergite]]. [[Ventral]]ly, somites 3 to 7 are armored with matching plates called [[Sternum (arthropod anatomy)|sternites]]. The ventral side of somite 1 has a pair of genital [[Operculum (animal)|opercula]] covering the [[gonopore]]. Sternite 2 forms the basal plate bearing the [[Pecten (biology)|pectines]],{{sfn|Polis|1990|pp=13–14}} which function as sensory organs.<ref name="Knowlton Gaffin 2011">{{cite journal |last1=Knowlton |first1=Elizabeth D. |last2=Gaffin |first2=Douglas D. |title=Functionally Redundant Peg Sensilla on the Scorpion Pecten |journal=Journal of Comparative Physiology A |publisher=Springer |volume=197 |issue=9 |year=2011 |issn=0340-7594 |doi=10.1007/s00359-011-0650-9 |pages=895–902|pmid=21647695 |s2cid=22123929 }}</ref>
+The mesosoma or preabdomen is the broad part of the opisthosoma.{{sfn|Polis|1990|pp=10–11}} In the early stages of embryonic development the mesosoma consist of eight segments, but the first segment disappear before birth, so the mesosoma in scorpions actually consist of segments 2-8.<ref>{{Cite journal|title=The ultrastructure of book lung development in the bark scorpion Centruroides gracilis (Scorpiones: Buthidae) - PMC|year=2011 |pmc=3199777 |last1=Farley |first1=R. D. |journal=Frontiers in Zoology |volume=8 |page=18 |doi=10.1186/1742-9994-8-18 |pmid=21791110 |doi-access=free }}</ref><ref>{{Cite web|url=http://www.sharmalabuw.org/uploads/1/3/6/1/13619635/schwager_et_al._2015.pdf|title=Chelicerata - Prashant P. Sharma}}</ref><ref>{{Cite web|url=http://www.science.marshall.edu/fet/euscorpius/p2011_120.pdf|title=Euscorpius - Marshall University}}</ref> These [[anterior]] seven [[somite]]s (segments) of the opisthosoma are each covered [[Anatomical terms of location|dorsally]] by a sclerotized plate called the [[tergite]]. [[Ventral]]ly, somites 3 to 7 are armored with matching plates called [[sternites]]. The ventral side of somite 1 has a pair of genital [[Operculum (animal)|opercula]] covering the [[gonopore]]. Sternite 2 forms the basal plate bearing the [[pectines]],{{sfn|Polis|1990|pp=13–14}} which function as sensory organs.<ref name="Knowlton Gaffin 2011">{{cite journal |last1=Knowlton |first1=Elizabeth D. |last2=Gaffin |first2=Douglas D. |title=Functionally Redundant Peg Sensilla on the Scorpion Pecten |journal=Journal of Comparative Physiology A |publisher=Springer |volume=197 |issue=9 |year=2011 |issn=0340-7594 |doi=10.1007/s00359-011-0650-9 |pages=895–902|pmid=21647695 |s2cid=22123929 }}</ref>
 The next four somites, 3 to 6, all bear pairs of [[Spiracle (arthropods)|spiracles]]. They serve as openings for the scorpion's respiratory organs, known as [[book lungs]]. The spiracle openings may be slits, circular, elliptical or oval according to the species.{{sfn|Polis|1990|p=15}}<ref name="Wanninger 2015">{{cite book |last=Wanninger |first=Andreas |title=Evolutionary Developmental Biology of Invertebrates 3: Ecdysozoa I: Non-Tetraconata |url=https://books.google.com/books?id=rxZcCgAAQBAJ&pg=PA105 |year=2015 |publisher=Springer |isbn=978-3-7091-1865-8 |page=105 |access-date=11 September 2017 |archive-date=5 August 2020 |archive-url=https://web.archive.org/web/20200805003251/https://books.google.com/books?id=rxZcCgAAQBAJ&pg=PA105 |url-status=live }}</ref> There are thus four pairs of book lungs; each consists of some 140 to 150 thin [[lamella (surface anatomy)|lamellae]] filled with air inside a pulmonary chamber, connected on the ventral side to an atrial chamber which opens into a spiracle. Bristles hold the lamellae apart. A muscle opens the spiracle and widens the atrial chamber; dorsoventral muscles contract to compress the pulmonary chamber, forcing air out, and relax to allow the chamber to refill.{{sfn|Polis|1990|pp=42–44}} The 7th and last somite does not bear appendages or any other significant external structures.{{sfn|Polis|1990|p=15}}
@@ Line 189: / Line 192: @@
 The "tail" or metasoma consists of five segments and the [[telson]], which is not strictly a segment. The five segments are merely body rings; they lack apparent sterna or terga, and become larger distally. These segments have keels, setae and bristles which may be used for taxonomic classification. The anus is at the distal and ventral end of the last segment, and is encircled by four anal papillae and the anal arch.{{sfn|Polis|1990|p=15}} The tails of some species contain light receptors.<ref name="Chakravarthy Sridhara 2016"/>
-The telson includes the [[vesicle (biology and chemistry)|vesicle]], which contains a symmetrical pair of [[venom]] glands. Externally it bears the curved stinger, the hypodermic aculeus, equipped with sensory hairs. Each of the venom glands has its own duct to convey its secretion along the aculeus from the bulb of the gland to immediately near of the tip, where each of the paired ducts has its own venom pore.<ref name="Yigit Benli 2010">{{cite journal |last1=Yigit |first1=N. |last2=Benli |first2=M. |title=Fine Structural Analysis of the Stinger in Venom Apparatus of the Scorpion ''Euscorpius mingrelicus'' (Scorpiones: Euscorpiidae) |journal=Journal of Venomous Animals and Toxins Including Tropical Diseases |date=2010 |volume=16 |issue=1 |pages=76–86 |doi=10.1590/s1678-91992010005000003 |doi-access=free}}</ref> An extrinsic muscle system in the tail moves it forward and propels and penetrates with the aculeus, while an intrinsic muscle system attached to the glands pumps venom through the stinger into the intended victim.{{sfn|Stockmann|2015|p=30}} The stinger contains [[metalloprotein]]s with zinc, hardening the tip.<ref>{{cite book |last=Schofield |first=R. M. S. |year=2001 |chapter=Metals in cuticular structures |title=Scorpion Biology and Research |editor1=Brownell, P. H. |editor2=Polis, G. A. |pages=234–256 |location=New York |publisher=Oxford University Press |isbn=978-0195084344}}</ref> The optimal stinging angle is around 30 degrees relative to the tip.<ref>{{cite journal |last1=van der Meijden |first1=Arie |last2=Kleinteich |first2=Thomas |title=A biomechanical view on stinger diversity in scorpions |journal=Journal of Anatomy |date=April 2017 |volume=230 |issue=4 |pages=497–509 |doi=10.1111/joa.12582|pmid=28028798 |pmc=5345679 |doi-access=free }}</ref><!-- +chitin fibril https://www.researchgate.net/publication/282399934_Study_of_biomechanical_anatomical_and_physiological_properties_of_scorpion_stingers_for_developing_biomimetic_materials -->
+The telson includes the [[vesicle (biology and chemistry)|vesicle]], which contains a symmetrical pair of [[venom gland]]s. Externally it bears the curved stinger, the hypodermic aculeus, equipped with sensory hairs. Each of the venom glands has its own duct to convey its secretion along the aculeus from the bulb of the gland to immediately near of the tip, where each of the paired ducts has its own venom pore.<ref name="Yigit Benli 2010">{{cite journal |last1=Yigit |first1=N. |last2=Benli |first2=M. |title=Fine Structural Analysis of the Stinger in Venom Apparatus of the Scorpion ''Euscorpius mingrelicus'' (Scorpiones: Euscorpiidae) |journal=Journal of Venomous Animals and Toxins Including Tropical Diseases |date=2010 |volume=16 |issue=1 |pages=76–86 |doi=10.1590/s1678-91992010005000003 |doi-access=free}}</ref> An extrinsic muscle system in the tail moves it forward and propels and penetrates with the aculeus, while an intrinsic muscle system attached to the glands pumps venom through the stinger into the intended victim.{{sfn|Stockmann|2015|p=30}} The stinger contains [[metalloprotein]]s with zinc, hardening the tip.<ref>{{cite book |last=Schofield |first=R. M. S. |year=2001 |chapter=Metals in cuticular structures |title=Scorpion Biology and Research |editor1=Brownell, P. H. |editor2=Polis, G. A. |pages=234–256 |location=New York |publisher=Oxford University Press |isbn=978-0195084344}}</ref> The optimal stinging angle is around 30 degrees relative to the tip.<ref>{{cite journal |last1=van der Meijden |first1=Arie |last2=Kleinteich |first2=Thomas |title=A biomechanical view on stinger diversity in scorpions |journal=Journal of Anatomy |date=April 2017 |volume=230 |issue=4 |pages=497–509 |doi=10.1111/joa.12582|pmid=28028798 |pmc=5345679 |doi-access=free }}</ref><!-- +chitin fibril https://www.researchgate.net/publication/282399934_Study_of_biomechanical_anatomical_and_physiological_properties_of_scorpion_stingers_for_developing_biomimetic_materials -->
 ==Biology==
@@ Line 196: / Line 199: @@
 Most scorpion species are [[nocturnal]] or [[crepuscular]], finding shelter during the day in burrows, cracks in rocks and tree bark.{{sfn|Stockmann|2015|pp=40–41}} Many species dig a shelter underneath stones a few centimeters long. Some may use burrows made by other animals including spiders, reptiles and small mammals. Other species dig their own burrows which vary in complexity and depth. ''[[Hadrurus]]'' species dig burrows as over {{convert|2|m|abbr=on}} deep. Digging is done using the mouth parts, claws and legs. In several species, particularly of the family Buthidae, individuals may gather in the same shelter; bark scorpions may aggregate up to 30 individuals. In some species, families of females and young sometimes aggregate.{{sfn|Stockmann|Ythier|2010|pp=146, 153–154}}
-Scorpions prefer areas where the temperature remains in the range of {{convert|11|–|40|C|F}}, but may survive temperatures from well below freezing to desert heat.<ref>{{cite journal |last=Hadley |first=Neil F. |title=Water Relations of the Desert Scorpion, ''Hadrurus arizonensis'' |journal=[[Journal of Experimental Biology]] |year=1970 |volume=53 |issue=3 |pages=547–558 |doi=10.1242/jeb.53.3.547 |url=https://journals.biologists.com/jeb/article/53/3/547/21677/Water-Relations-of-the-Desert-Scorpion-Hadrurus |pmid=5487163 |access-date=30 December 2006 |archive-date=6 April 2008 |archive-url=https://web.archive.org/web/20080406195500/http://jeb.biologists.org/cgi/reprint/53/3/547.pdf |url-status=live }}</ref><ref>{{cite journal |last1=Hoshino |first1=K. |last2=Moura |first2=A. T. V. |last3=De Paula |first3=H. M. G. |title=Selection of Environmental Temperature by the Yellow Scorpion ''Tityus serrulatus'' Lutz & Mello, 1922 (Scorpiones, Buthidae) |journal=Journal of Venomous Animals and Toxins Including Tropical Diseases |year=2006 |volume=12 |issue=1 |pages=59–66 |doi=10.1590/S1678-91992006000100005 |doi-access=free }}</ref> Scorpions can withstand intense heat: ''[[Leiurus quinquestriatus]]'', ''Scorpio maurus''<!--linked above--> and ''[[Hadrurus arizonensis]]'' can live in temperatures of {{convert|45|–|50|C|F|0|abbr=on}} if they are sufficiently hydrated. Desert species must deal with the extreme changes in temperature from day to night or between seasons; ''[[Pectinibuthus birulai]]'' lives in a temperature range of {{convert|-30|–|50|C|F|0|abbr=on}}. Scorpions that live outside deserts prefer lower temperatures. The ability to resist cold may be related to the increase in the sugar [[trehalose]] when the temperature drops. Some species [[hibernate]].{{sfn|Stockmann|2015|pp=42–43}} Scorpions appear to have resistance to [[ionizing radiation]]. This was discovered in the early 1960s when scorpions were found to be among the few animals to survive nuclear tests at [[Reggane]], Algeria.{{sfn|Stockmann|Ythier|2010|p=157}}
+Scorpions prefer areas where the temperature remains in the range of {{convert|11|–|40|C|F}}, but may survive temperatures from well below freezing to desert heat.<ref>{{cite journal |last=Hadley |first=Neil F. |title=Water Relations of the Desert Scorpion, ''Hadrurus arizonensis'' |journal=[[Journal of Experimental Biology]] |year=1970 |volume=53 |issue=3 |pages=547–558 |doi=10.1242/jeb.53.3.547 |url=https://journals.biologists.com/jeb/article/53/3/547/21677/Water-Relations-of-the-Desert-Scorpion-Hadrurus |pmid=5487163 |access-date=30 December 2006 |archive-date=6 April 2008 |archive-url=https://web.archive.org/web/20080406195500/http://jeb.biologists.org/cgi/reprint/53/3/547.pdf |url-status=live }}</ref><ref>{{cite journal |last1=Hoshino |first1=K. |last2=Moura |first2=A. T. V. |last3=De Paula |first3=H. M. G. |title=Selection of Environmental Temperature by the Yellow Scorpion ''Tityus serrulatus'' Lutz & Mello, 1922 (Scorpiones, Buthidae) |journal=Journal of Venomous Animals and Toxins Including Tropical Diseases |year=2006 |volume=12 |issue=1 |pages=59–66 |doi=10.1590/S1678-91992006000100005 |doi-access=free |hdl=11449/68851 |hdl-access=free }}</ref> Scorpions can withstand intense heat: ''[[Leiurus quinquestriatus]]'', ''Scorpio maurus''<!--linked above--> and ''[[Hadrurus arizonensis]]'' can live in temperatures of {{convert|45|–|50|C|F|0|abbr=on}} if they are sufficiently hydrated. Desert species must deal with the extreme changes in temperature from day to night or between seasons; ''[[Pectinibuthus birulai]]'' lives in a temperature range of {{convert|-30|–|50|C|F|0|abbr=on}}. Scorpions that live outside deserts prefer lower temperatures. The ability to resist cold may be related to the increase in the sugar [[trehalose]] when the temperature drops. Some species [[hibernate]].{{sfn|Stockmann|2015|pp=42–43}} Scorpions appear to have resistance to [[ionizing radiation]]. This was discovered in the early 1960s when scorpions were found to be among the few animals to survive nuclear tests at [[Reggane]], Algeria.{{sfn|Stockmann|Ythier|2010|p=157}}
 Desert scorpions have several adaptations for water conservation. They excrete insoluble compounds such as [[xanthine]], [[guanine]], and [[uric acid]], not requiring water for their removal from the body. Guanine is the main component and maximizes the amount of nitrogen excreted. A scorpion's cuticle holds in moisture via [[lipid]]s and waxes from epidermal glands, and protects against [[ultraviolet radiation]]. Even when dehydrated, a scorpion can tolerate high [[osmotic pressure]] in its blood.<ref>{{cite book |last=Cowles |first=Jillian |title=Amazing Arachnids |publisher=Princeton University Press |year=2018 |page=33 |isbn=978-0-691-17658-1}}</ref> Desert scorpions get most of their moisture from the food they eat but some can absorb water from the humid soil. Species that live in denser vegetation and in more moderate temperatures will drink water on plants and in puddles.{{sfn|Stockmann|Ythier|2010|p=156}}
 [[File:Venom Squirting in Scorpions.svg|thumb|upright=0.9|A few scorpions squirt [[venom]] to deter predators.]]
-A scorpion uses its stinger both for killing prey and defense. Some species make direct, quick strikes with their tails while others make slower, more circular strikes which can more easily return the stinger to a position where it can strike again. ''Leiurus quinquestriatus'' can whip its tail at a speed of up to {{convert|128|cm/s|in/s|abbr=on}}<!--Coelho et al, table 3--> in a defensive strike.<ref>{{cite journal |last1=Coelho |first1=P. |last2=Kaliontzopoulou |first2=A. |last3=Rasko |first3=M. |last4=van der Meijden |first4=A. |year=2017 |title=A 'Striking' Relationship: Scorpion Defensive Behaviour and its Relation to Morphology and Performance |journal=Functional Ecology |volume=31|issue=7 |pages=1390–1404 |doi=10.1111/1365-2435.12855|doi-access=free }}</ref>
+A scorpion uses its stinger both for killing prey and defense. Some species make direct, quick strikes with their tails while others make slower, more circular strikes which can more easily return the stinger to a position where it can strike again. ''Leiurus quinquestriatus'' can whip its tail at a speed of up to {{convert|128|cm/s|in/s|abbr=on}}<!--Coelho et al, table 3--> in a defensive strike.<ref>{{cite journal |last1=Coelho |first1=P. |last2=Kaliontzopoulou |first2=A. |last3=Rasko |first3=M. |last4=van der Meijden |first4=A. |year=2017 |title=A 'Striking' Relationship: Scorpion Defensive Behaviour and its Relation to Morphology and Performance |journal=Functional Ecology |volume=31|issue=7 |pages=1390–1404 |doi=10.1111/1365-2435.12855|doi-access=free |bibcode=2017FuEco..31.1390C }}</ref>
 ===Mortality and defense===
@@ Line 210: / Line 213: @@
 [[File:Scorpion eating solifugid (Marshal Hedin).jpg|thumb|right<!--left-->|Scorpion feeding on a [[solifugid]]]]
-Scorpions generally prey on insects, particularly [[grasshopper]]s, [[Cricket (insect)|cricket]]s, [[termite]]s, [[beetle]]s and [[wasp]]s. Other prey include spiders, [[solifugid]]s, [[woodlice]] and even small [[vertebrate]]s including lizards, snakes and mammals. Species with large claws may prey on earthworms and mollusks. The majority of species are opportunistic and consume a variety of prey though some may be highly specialized; ''[[Isometroides vescus]]'' specializes on burrowing spiders. Prey size depends on the size of the species. Several scorpion species are [[sit-and-wait predator]]s, which involves them waiting for prey at or near the entrance to their burrow. Others actively seek them out. Scorpions detect their prey with [[Mechanoreceptor|mechanoreceptive]] and [[Chemoreceptor|chemoreceptive]] hairs on their bodies and capture them with their claws. Small animals are merely killed with the claws, particularly by large-clawed species. Larger and more aggressive prey is given a sting.{{sfn|Stockmann|2015|pp=35–38}}<ref>{{cite web |last=Murray |first=Melissa |title=Scorpions |publisher=Australian Museum |url=https://australian.museum/learn/animals/spiders/scorpions/#:~:text=Scorpions%20are%20nocturnal%20hunters%2C%20feeding,and%20often%20occupying%20their%20burrows |date=3 December 2020 |access-date=13 December 2020 |archive-date=8 November 2020 |archive-url=https://web.archive.org/web/20201108102328/https://australian.museum/learn/animals/spiders/scorpions/#:~:text=Scorpions%20are%20nocturnal%20hunters%2C%20feeding,and%20often%20occupying%20their%20burrows |url-status=live }}</ref>
+Scorpions generally prey on insects, particularly [[grasshopper]]s, [[Cricket (insect)|cricket]]s, [[termite]]s, [[beetle]]s and [[wasp]]s. Other prey include spiders, [[solifugid]]s, [[woodlice]] and even small [[vertebrate]]s including lizards, snakes and mammals. Species with large claws may prey on earthworms and mollusks. The majority of species are opportunistic and consume a variety of prey though some may be highly specialized; ''[[Isometroides vescus]]'' specializes on burrowing spiders. Prey size depends on the size of the species. Several scorpion species are [[sit-and-wait predator]]s, which involves them waiting for prey at or near the entrance to their burrow. Others actively seek them out. Scorpions detect their prey with [[mechanoreceptive]] and [[chemoreceptive]] hairs on their bodies and capture them with their claws. Small animals are merely killed with the claws, particularly by large-clawed species. Larger and more aggressive prey is given a sting.{{sfn|Stockmann|2015|pp=35–38}}<ref>{{cite web |last=Murray |first=Melissa |title=Scorpions |publisher=Australian Museum |url=https://australian.museum/learn/animals/spiders/scorpions/#:~:text=Scorpions%20are%20nocturnal%20hunters%2C%20feeding,and%20often%20occupying%20their%20burrows |date=3 December 2020 |access-date=13 December 2020 |archive-date=8 November 2020 |archive-url=https://web.archive.org/web/20201108102328/https://australian.museum/learn/animals/spiders/scorpions/#:~:text=Scorpions%20are%20nocturnal%20hunters%2C%20feeding,and%20often%20occupying%20their%20burrows |url-status=live }}</ref>
 Scorpions, like other arachnids, digest their food externally. The chelicerae, which are very sharp, are used to pull small amounts of food off the prey item into a pre-oral cavity below the chelicerae and carapace. The digestive juices from the gut are egested onto the food, and the digested food is then sucked into the gut in liquid form. Any solid indigestible matter (such as [[exoskeleton]] fragments) is trapped by [[seta]]e in the pre-oral cavity and ejected. The sucked-in food is pumped into the midgut by the [[pharynx]], where it is further digested. The waste passes through the hindgut and out of the anus. Scorpions can consume large amounts of food during one meal. They have an efficient food storage organ and a very low [[metabolic rate]], and a relatively inactive lifestyle. This enables some to survive six to twelve months of starvation.{{sfn|Polis|1990|pp=296–298}}
@@ Line 226: / Line 229: @@
 [[File:scorpionwithyoung.JPG|thumb|right<!--left-->|''[[Compsobuthus werneri]]'' female with [[Offspring|young]]]]
-[[Gestation]] in scorpions can last for over a year in some species.{{sfn|Polis|1990|p=161}} They have two types of [[embryonic development]]; apoikogenic and katoikogenic. In the apoikogenic system, which is mainly found in the Buthidae, embryos develop in yolk-rich eggs inside [[Ovarian follicle|follicles]]. The katoikogenic system is documented in Hemiscorpiidae, Scorpionidae and Diplocentridae, and involves the embryos developing in a [[diverticulum]] which has a teat-like structure for them to feed though.<ref>{{cite journal |last=Warburg |first=M. R. |year=2010 |title=Reproductive System of Female Scorpion: A Partial Review |journal=The Anatomical Record |volume=293 |issue=10 |pages=1738–1754 |doi=10.1002/ar.21219 |pmid=20687160 |s2cid=25391120 |doi-access=free}}</ref> Unlike the majority of arachnids, which are [[oviparous]], hatching from eggs, scorpions seem to be universally [[viviparous]], with live births.<ref name="Warburg">{{cite journal |last1=Warburg |first1=Michael R. |title=Pre- and Post-parturial Aspects of Scorpion Reproduction: a Review |journal=European Journal of Entomology |date=5 April 2012 |volume=109 |issue=2 |pages=139–146 |doi=10.14411/eje.2012.018 |doi-access=free }}</ref> They are unusual among terrestrial arthropods in the amount of care a female gives to her offspring.{{sfn|Polis|1990|p=6}} The size of a brood varies by species, from 3 to over 100.<ref name="Scharff"/> The body size of scorpions is not correlated either with brood size or with life cycle length.<ref>{{cite journal |last=Monge-Nájera |first=J. |year=2019 |title=Scorpion Body Size, Litter Characteristics, and Duration of the Life Cycle (Scorpiones) |journal=Cuadernos de Investigación UNED |volume=11 |issue=2 |pages=101–104}}</ref>
+[[Gestation]] in scorpions can last for over a year in some species.{{sfn|Polis|1990|p=161}} They have two types of [[embryonic development]]; apoikogenic and katoikogenic. In the apoikogenic system, which is mainly found in the Buthidae, embryos develop in yolk-rich eggs inside [[Ovarian follicle|follicles]]. The katoikogenic system is documented in Hemiscorpiidae, Scorpionidae and Diplocentridae, and involves the embryos developing in a [[diverticulum]] which has a teat-like structure for them to feed through.<ref>{{cite journal |last=Warburg |first=M. R. |year=2010 |title=Reproductive System of Female Scorpion: A Partial Review |journal=The Anatomical Record |volume=293 |issue=10 |pages=1738–1754 |doi=10.1002/ar.21219 |pmid=20687160 |s2cid=25391120 |doi-access=free}}</ref> Unlike the majority of arachnids, which are [[oviparous]], hatching from eggs, scorpions seem to be universally [[viviparous]], with live births.<ref name="Warburg">{{cite journal |last1=Warburg |first1=Michael R. |title=Pre- and Post-parturial Aspects of Scorpion Reproduction: a Review |journal=European Journal of Entomology |date=5 April 2012 |volume=109 |issue=2 |pages=139–146 |doi=10.14411/eje.2012.018 |doi-access=free }}</ref> They are unusual among terrestrial arthropods in the amount of care a female gives to her offspring.{{sfn|Polis|1990|p=6}} The size of a brood varies by species, from 3 to over 100.<ref name="Scharff"/> The body size of scorpions is not correlated either with brood size or with life cycle length.<ref>{{cite journal |last=Monge-Nájera |first=J. |year=2019 |title=Scorpion Body Size, Litter Characteristics, and Duration of the Life Cycle (Scorpiones) |journal=Cuadernos de Investigación UNED |volume=11 |issue=2 |pages=101–104}}</ref>
 Before giving birth, the female elevates the front of her body and positions her pedipalps and front legs under her to catch the young ("birth basket"). The young emerge one by one from the genital opercula, expel the embryonic membrane, if any, and are placed on the mother's back where they remain until they have gone through at least one [[ecdysis|molt]]. The period before the first molt is called the pro-juvenile stage; the young are unable to feed or sting, but have [[Sucker (zoology)|suckers]] on their tarsi, used to hold on to their mother. This period lasts 5 to 25 days, depending on the species. The brood molt for the first time simultaneously in a process that lasts 6 to 8 hours, marking the beginning of the juvenile stage.<ref name="Scharff">{{cite book |last=Lourenço |first=Wilson R. |author-link=Wilson R. Lourenço |title=European Arachnology |chapter=Reproduction in Scorpions, with Special Reference to Parthenogenesis |year=2000 |editor1-first=S. |editor1-last=Toft |editor2-first=N. |editor2-last=Scharff |publisher=Aarhus University Press |isbn=978-877934-0015 |pages=74–76 |chapter-url=http://www.european-arachnology.org/wdp/wp-content/uploads/2015/08/071-085_Lourenco.pdf |access-date=14 December 2020 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031223/http://www.european-arachnology.org/wdp/wp-content/uploads/2015/08/071-085_Lourenco.pdf |url-status=live }}</ref>
@@ Line 235: / Line 238: @@
 {{multiple image|image1=Sorpion Under Blacklight edit.jpg|caption1=This black [[emperor scorpion]] (''Pandinus imperator'') [[fluoresce]]s light blue.|width1=200|image2=Scorpion with babies in UV.jpg|caption2=The mother glows bright teal, the young a dull grey.|width2=113}}
-Scorpions glow a vibrant blue-green when exposed to certain wavelengths ranges of [[ultraviolet]] light such as that produced by a [[black light]], due to [[fluorescent]] chemicals such as [[beta-carboline]] in the cuticle. Accordingly, a hand-held ultraviolet lamp has long been a standard tool for nocturnal field surveys of these animals. Fluorescence occurs as a result of [[sclerotization]] and increases in intensity with each successive instar.<ref name="Stachel 1999 531–539">{{cite journal |first1=Shawn J. |last1=Stachel |first2=Scott A. |last2=Stockwell |first3=David L. |last3=Van Vranken |date=August 1999 |title=The Fluorescence of Scorpions and Cataractogenesis |journal=Chemistry & Biology |volume=6 |pages=531–539 |doi=10.1016/S1074-5521(99)80085-4 |pmid=10421760 |issue=8 |doi-access=free }}</ref> This fluorescence may have an active role in the scorpion's ability to detect light.<ref>{{cite journal |last1=Gaffinr |first1=Douglas D. |last2=Bumm |first2=Lloyd A. |last3=Taylor |first3=Matthew S. |last4=Popokina |first4=Nataliya V. |last5=Mann |first5=Shivani |year=2012 |title=Scorpion Fluorescence and Reaction to Light |journal=[[Animal Behaviour (journal)|Animal Behaviour]] |issue=2 |doi=10.1016/j.anbehav.2011.11.014 |volume=83 |pages=429–436 |s2cid=17041988 |url=https://zenodo.org/record/895450 |access-date=18 May 2018 |archive-date=6 November 2020 |archive-url=https://web.archive.org/web/20201106153700/https://zenodo.org/record/895450 |url-status=live }}</ref>
+Scorpions glow a vibrant blue-green when exposed to certain wavelengths of [[ultraviolet]] light, such as that produced by a [[black light]], due to [[fluorescent]] chemicals such as [[beta-carboline]] in the cuticle. Accordingly, a hand-held ultraviolet lamp has long been a standard tool for nocturnal field surveys of these animals. Fluorescence occurs as a result of [[sclerotization]] and increases in intensity with each successive instar.<ref name="Stachel 1999 531–539">{{cite journal |first1=Shawn J. |last1=Stachel |first2=Scott A. |last2=Stockwell |first3=David L. |last3=Van Vranken |date=August 1999 |title=The Fluorescence of Scorpions and Cataractogenesis |journal=Chemistry & Biology |volume=6 |pages=531–539 |doi=10.1016/S1074-5521(99)80085-4 |pmid=10421760 |issue=8 |doi-access=free }}</ref> This fluorescence may have an active role in the scorpion's ability to detect light.<ref>{{cite journal |last1=Gaffinr |first1=Douglas D. |last2=Bumm |first2=Lloyd A. |last3=Taylor |first3=Matthew S. |last4=Popokina |first4=Nataliya V. |last5=Mann |first5=Shivani |year=2012 |title=Scorpion Fluorescence and Reaction to Light |journal=[[Animal Behaviour (journal)|Animal Behaviour]] |issue=2 |doi=10.1016/j.anbehav.2011.11.014 |volume=83 |pages=429–436 |s2cid=17041988 |url=https://zenodo.org/record/895450 |access-date=18 May 2018 |archive-date=6 November 2020 |archive-url=https://web.archive.org/web/20201106153700/https://zenodo.org/record/895450 |url-status=live }}</ref>
 {{anchor|In culture}}
@@ Line 246: / Line 249: @@
 [[File:Bbasgen-scorpion-front.jpg|thumb|right|upright|[[Arizona bark scorpion]], one of the few species whose venom is deadly to humans]]
-Scorpion venom serves to kill or paralyze prey rapidly. The [[Scorpion sting|sting]]s of many species are uncomfortable, but only 25 species have venom that is deadly to humans. Those species belong to the family Buthidae, including ''Leiurus quinquestriatus'', ''Hottentotta'' spp., ''[[Centruroides]]'' spp., and ''[[Androctonus]]'' spp.<ref name="Earthlife"/> People with [[Allergy|allergies]] are especially at risk;<ref name="niosh">{{cite web |url=https://www.cdc.gov/niosh/topics/insects/scorpions.html |title=Insects and Scorpions |publisher=[[NIOSH]] |date=1 July 2016 |access-date=15 July 2016 |archive-date=29 July 2016 |archive-url=https://web.archive.org/web/20160729121849/http://www.cdc.gov/niosh/topics/insects/scorpions.html |url-status=live }}</ref> otherwise, [[first aid]] is [[Symptomatic treatment|symptomatic]], with [[Analgesic|analgesia]]. [[Hypertensive emergency|Cases of very high blood pressure]] are treated with medications that [[anxiolytic|relieve anxiety]] and [[Vasodilation|relax the blood vessels]].<ref>{{cite book |last1=Buma |first1=Adriaan Hopperus |last2=Burris |first2=David G. |last3=Hawley |first3=Alan |last4=Ryan |first4=James M. |last5=Mahoney |first5=Peter F. |title=Conflict and Catastrophe Medicine: A Practical Guide |year=2009 |edition=2nd |publisher=Springer |isbn=978-1-84800-351-4 |chapter=Scorpion sting |page=518 |chapter-url=https://books.google.com/books?id=Q-kk_TX9Ji0C&pg=PA518 |access-date=5 March 2016 |archive-date=25 November 2016 |archive-url=https://web.archive.org/web/20161125152534/https://books.google.com/books?id=Q-kk_TX9Ji0C&pg=PA518 |url-status=live }}</ref><ref name=mayo>{{cite web |title=Diseases and Conditions – Scorpion Stings |url=http://www.mayoclinic.org/diseases-conditions/scorpion-stings/basics/definition/con-20033894 |publisher=[[Mayo Clinic]] |access-date=3 July 2015 |archive-date=4 October 2016 |archive-url=https://web.archive.org/web/20161004042604/http://www.mayoclinic.org/diseases-conditions/scorpion-stings/basics/definition/con-20033894 |url-status=live }}</ref> Scorpion envenomation with high morbidity and mortality is usually due to either excessive [[Autonomic nervous system|autonomic activity]] and cardiovascular toxic effects, or neuromuscular toxic effects. [[Antivenom]] is the specific treatment for scorpion envenomation combined with supportive measures including vasodilators in patients with cardiovascular toxic effects, and [[benzodiazepine]]s when there is neuromuscular involvement. Although rare, severe [[hypersensitivity reaction]]s including [[anaphylaxis]] to scorpion antivenin are possible.<ref name="10.4103/0972-5229.164807">{{cite journal |last1=Bhoite |first1=R. R. |last2=Bhoite |first2=G. R. |last3=Bagdure |first3=D. N. |last4=Bawaskar |first4=H. S. |title=Anaphylaxis to Scorpion Antivenin and its Management Following Envenomation by Indian Red Scorpion, Mesobuthus tamulus |journal=[[Indian Journal of Critical Care Medicine]] |volume=19 |issue=9 |pages=547–549 |year=2015 |doi=10.4103/0972-5229.164807 |pmc=4578200 |pmid=26430342}}</ref>
+Scorpion venom serves to kill or paralyze prey rapidly. The [[Scorpion sting|sting]]s of many species are uncomfortable, but only 25 species have venom that is deadly to humans. Those species belong to the family Buthidae, including ''Leiurus quinquestriatus'', ''Hottentotta'' spp., ''[[Centruroides]]'' spp., and ''[[Androctonus]]'' spp.<ref name="Earthlife"/> People with [[allergies]] are especially at risk;<ref name="niosh">{{cite web |url=https://www.cdc.gov/niosh/topics/insects/scorpions.html |title=Insects and Scorpions |publisher=[[NIOSH]] |date=1 July 2016 |access-date=15 July 2016 |archive-date=29 July 2016 |archive-url=https://web.archive.org/web/20160729121849/http://www.cdc.gov/niosh/topics/insects/scorpions.html |url-status=live }}</ref> otherwise, [[first aid]] is [[Symptomatic treatment|symptomatic]], with [[Analgesic|analgesia]]. [[Hypertensive emergency|Cases of very high blood pressure]] are treated with medications that [[anxiolytic|relieve anxiety]] and [[Vasodilation|relax the blood vessels]].<ref>{{cite book |last1=Buma |first1=Adriaan Hopperus |last2=Burris |first2=David G. |last3=Hawley |first3=Alan |last4=Ryan |first4=James M. |last5=Mahoney |first5=Peter F. |title=Conflict and Catastrophe Medicine: A Practical Guide |year=2009 |edition=2nd |publisher=Springer |isbn=978-1-84800-351-4 |chapter=Scorpion sting |page=518 |chapter-url=https://books.google.com/books?id=Q-kk_TX9Ji0C&pg=PA518 |access-date=5 March 2016 |archive-date=25 November 2016 |archive-url=https://web.archive.org/web/20161125152534/https://books.google.com/books?id=Q-kk_TX9Ji0C&pg=PA518 |url-status=live }}</ref><ref name=mayo>{{cite web |title=Diseases and Conditions – Scorpion Stings |url=http://www.mayoclinic.org/diseases-conditions/scorpion-stings/basics/definition/con-20033894 |publisher=[[Mayo Clinic]] |access-date=3 July 2015 |archive-date=4 October 2016 |archive-url=https://web.archive.org/web/20161004042604/http://www.mayoclinic.org/diseases-conditions/scorpion-stings/basics/definition/con-20033894 |url-status=live }}</ref> Scorpion envenomation with high morbidity and mortality is usually due to either excessive [[Autonomic nervous system|autonomic activity]] and cardiovascular toxic effects, or neuromuscular toxic effects. [[Antivenom]] is the specific treatment for scorpion envenomation combined with supportive measures including vasodilators in patients with cardiovascular toxic effects, and [[benzodiazepine]]s when there is neuromuscular involvement. Although rare, severe [[hypersensitivity reaction]]s including [[anaphylaxis]] to scorpion antivenin are possible.<ref name="10.4103/0972-5229.164807">{{cite journal |last1=Bhoite |first1=R. R. |last2=Bhoite |first2=G. R. |last3=Bagdure |first3=D. N. |last4=Bawaskar |first4=H. S. |title=Anaphylaxis to Scorpion Antivenin and its Management Following Envenomation by Indian Red Scorpion, Mesobuthus tamulus |journal=[[Indian Journal of Critical Care Medicine]] |volume=19 |issue=9 |pages=547–549 |year=2015 |doi=10.4103/0972-5229.164807 |pmc=4578200 |pmid=26430342 |doi-access=free }}</ref>
-Scorpion stings are a public health problem, particularly in the tropical and subtropical regions of the Americas, North Africa, the Middle East and India. Around 1.5 million scorpion envenomations occur each year with around 2,600 deaths.<ref name="Feola 2020">{{cite journal |last1=Feola |first1=A. |last2=Perrone |first2=M. A. |last3=Piscopo |first3=A. |last4=Casella |first4=F. |last5=Della Pietra |first5=B. |last6=Di Mizio |first6=G. |year=2020 |title=Autopsy Findings in Case of Fatal Scorpion Sting: A Systematic Review of the Literature |journal=Healthcare |volume=8 |issue=3 |page=325 |doi=10.3390/healthcare8030325|pmid=32899951 |pmc=7551928 |doi-access=free }}</ref>{{sfn|Stockmann|Ythier|2010|pp=163–164}}<ref name="Santos Silva 2016">{{cite journal |last1=Santos |first1=Maria S.V. |last2=Silva |first2=Cláudio G.L. |last3=Neto |first3=Basílio Silva |last4=Grangeiro Júnior |first4=Cícero R.P. |last5=Lopes |first5=Victor H.G. |last6=Teixeira Júnior |first6=Antônio G. |last7=Bezerra |first7=Deryk A. |last8=Luna |first8=João V.C.P. |last9=Cordeiro |first9=Josué B. |last10=Júnior |first10=Jucier Gonçalves |last11=Lima |first11=Marcos A.P. |title=Clinical and Epidemiological Aspects of Scorpionism in the World: A Systematic Review |journal=Wilderness & Environmental Medicine |volume=27 |issue=4 |year=2016 |pages=504–518 |issn=1080-6032 |doi=10.1016/j.wem.2016.08.003 |pmid=27912864 |url=https://www.wemjournal.org/article/S1080-6032(16)30214-9/pdf<!--not redundant to DOI--> |doi-access=free |access-date=8 December 2020 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031227/https://www.wemjournal.org/article/S1080-6032%2816%2930214-9/pdf |url-status=live }}</ref> Mexico is one of the most affected countries, with the highest [[biodiversity]] of scorpions in the world, some 200,000 envenomations per year and at least 300 deaths<!--more likely 1000-->.<ref name="Dehesa-DávilaPossani1994">{{cite journal |last1=Dehesa-Dávila |first1=Manuel |last2=Possani |first2=Lourival D. |title=Scorpionism and Serotherapy in Mexico |journal=Toxicon |volume=32 |issue=9 |year=1994 |pages=1015–1018 |issn=0041-0101 |doi=10.1016/0041-0101(94)90383-2|pmid=7801335 }}</ref><ref name="Santibáñez-LópezFrancke2015">{{cite journal |last1=Santibáñez-López |first1=Carlos |last2=Francke |first2=Oscar |last3=Ureta |first3=Carolina |last4=Possani |first4=Lourival |title=Scorpions from Mexico: From Species Diversity to Venom Complexity |journal=Toxins |volume=8 |issue=1 |year=2015 |pages=2 |issn=2072-6651 |doi=10.3390/toxins8010002|pmid=26712787 |pmc=4728524 |doi-access=free }}</ref>
+Scorpion stings are a public health problem, particularly in the tropical and subtropical regions of the Americas, North Africa, the Middle East and India. Around 1.5 million scorpion envenomations occur each year with around 2,600 deaths.<ref name="Feola 2020">{{cite journal |last1=Feola |first1=A. |last2=Perrone |first2=M. A. |last3=Piscopo |first3=A. |last4=Casella |first4=F. |last5=Della Pietra |first5=B. |last6=Di Mizio |first6=G. |year=2020 |title=Autopsy Findings in Case of Fatal Scorpion Sting: A Systematic Review of the Literature |journal=Healthcare |volume=8 |issue=3 |page=325 |doi=10.3390/healthcare8030325|pmid=32899951 |pmc=7551928 |doi-access=free }}</ref>{{sfn|Stockmann|Ythier|2010|pp=163–164}}<ref name="Santos Silva 2016">{{cite journal |last1=Santos |first1=Maria S.V. |last2=Silva |first2=Cláudio G.L. |last3=Neto |first3=Basílio Silva |last4=Grangeiro Júnior |first4=Cícero R.P. |last5=Lopes |first5=Victor H.G. |last6=Teixeira Júnior |first6=Antônio G. |last7=Bezerra |first7=Deryk A. |last8=Luna |first8=João V.C.P. |last9=Cordeiro |first9=Josué B. |last10=Júnior |first10=Jucier Gonçalves |last11=Lima |first11=Marcos A.P. |title=Clinical and Epidemiological Aspects of Scorpionism in the World: A Systematic Review |journal=Wilderness & Environmental Medicine |volume=27 |issue=4 |year=2016 |pages=504–518 |issn=1080-6032 |doi=10.1016/j.wem.2016.08.003 |pmid=27912864 |url=https://www.wemjournal.org/article/S1080-6032(16)30214-9/pdf<!--not redundant to DOI--> |doi-access=free |access-date=8 December 2020 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031227/https://www.wemjournal.org/article/S1080-6032%2816%2930214-9/pdf |url-status=live }}</ref> Mexico is one of the most affected countries, with the highest [[biodiversity]] of scorpions in the world, some 200,000 envenomations per year and at least 300 deaths<!--more likely 1000-->.<ref name="Dehesa-DávilaPossani1994">{{cite journal |last1=Dehesa-Dávila |first1=Manuel |last2=Possani |first2=Lourival D. |title=Scorpionism and Serotherapy in Mexico |journal=Toxicon |volume=32 |issue=9 |year=1994 |pages=1015–1018 |issn=0041-0101 |doi=10.1016/0041-0101(94)90383-2|pmid=7801335 |bibcode=1994Txcn...32.1015D }}</ref><ref name="Santibáñez-LópezFrancke2015">{{cite journal |last1=Santibáñez-López |first1=Carlos |last2=Francke |first2=Oscar |last3=Ureta |first3=Carolina |last4=Possani |first4=Lourival |title=Scorpions from Mexico: From Species Diversity to Venom Complexity |journal=Toxins |volume=8 |issue=1 |year=2015 |pages=2 |issn=2072-6651 |doi=10.3390/toxins8010002|pmid=26712787 |pmc=4728524 |doi-access=free }}</ref>
 Efforts are made to prevent envenomation and to control scorpion populations. Prevention encompasses personal activities such as checking shoes and clothes before putting them on, not walking in bare feet or sandals, and filling in holes and cracks where scorpions might nest. Street lighting reduces scorpion activity. Control may involve the use of [[insecticide]]s such as [[pyrethroids]], or gathering scorpions manually with the help of ultraviolet lights. Domestic predators of scorpions, such as chickens and turkeys, can help to reduce the risk to a household.<ref name="Feola 2020"/>{{sfn|Stockmann|Ythier|2010|pp=163–164}}
@@ Line 255: / Line 258: @@
 [[File:Chlorotoxin.png|thumb|upright|The [[deathstalker]]'s powerful venom contains the 36-[[amino acid]] peptide [[chlorotoxin]] ([[ribbon diagram]] shown). This blocks small-conductance [[chloride channel]]s, immobilizing its prey.<ref name="Ahmadi Knerr Argemi Bordon p=118"/>]]
-Scorpion venom is a mixture of neurotoxins; most of these are [[peptide]]s, chains of [[amino acid]]s.<ref name="Rodríguez 2013">{{cite book |last1=Rodríguez de la Vega |first1=Ricardo C. |last2=Vidal |first2=Nicolas |last3=Possani |first3=Lourival D. |chapter=Scorpion Peptides |editor-first=Abba J. |editor-last=Kastin |title=Handbook of Biologically Active Peptides |publisher=Academic Press |edition=2nd |year=2013 |isbn=978-0-12-385095-9 |pages=423–429 |doi=10.1016/B978-0-12-385095-9.00059-2}}</ref> Many of them interfere with [[membrane channel]]s that [[Sodium channel|transport sodium]], [[Potassium channel|potassium]], [[calcium channel|calcium]], or [[Chloride channel|chloride]] ions. These channels are essential for [[nerve conduction]], [[muscle contraction]] and many other biological processes. Some of these molecules may be useful in medical research and might lead to the development of new disease treatments. Among their potential therapeutic uses are as analgesic, [[anti-cancer]], [[antibacterial]], [[antifungal]], [[antiviral]], [[antiparasitic]], [[bradykinin]]-potentiating, and [[immunosuppressive]] drugs. As of 2020, no scorpion toxin-based drug is on sale, though [[chlorotoxin]] is being trialled for use against [[glioma]], a brain cancer.<ref name="Ahmadi Knerr Argemi Bordon p=118">{{cite journal |last1=Ahmadi |first1=Shirin |last2=Knerr |first2=Julius M. |last3=Argemi |first3=Lídia |last4=Bordon |first4=Karla C. F. |last5=Pucca |first5=Manuela B. |last6=Cerni |first6=Felipe A. |last7=Arantes |first7=Eliane C. |last8=Çalışkan |first8=Figen |last9=Laustsen |first9=Andreas H. |title=Scorpion Venom: Detriments and Benefits |journal=Biomedicines |publisher=MDPI AG |volume=8 |issue=5 |date=12 May 2020 |issn=2227-9059 |doi=10.3390/biomedicines8050118 |pmid=32408604 |pmc=7277529 |page=118|doi-access=free }}</ref>
+Scorpion venom is a mixture of neurotoxins; most of these are [[peptide]]s, chains of [[amino acid]]s.<ref name="Rodríguez 2013">{{cite book |last1=Rodríguez de la Vega |first1=Ricardo C. |last2=Vidal |first2=Nicolas |last3=Possani |first3=Lourival D. |chapter=Scorpion Peptides |editor-first=Abba J. |editor-last=Kastin |title=Handbook of Biologically Active Peptides |publisher=Academic Press |edition=2nd |year=2013 |isbn=978-0-12-385095-9 |pages=423–429 |doi=10.1016/B978-0-12-385095-9.00059-2}}</ref> Many of them interfere with [[membrane channel]]s that [[Sodium channel|transport sodium]], [[Potassium channel|potassium]], [[calcium channel|calcium]], or [[Chloride channel|chloride]] ions. These channels are essential for [[nerve conduction]], [[muscle contraction]] and many other biological processes. Some of these molecules may be useful in medical research and might lead to the development of new disease treatments. Among their potential therapeutic uses are as analgesic, [[anti-cancer]], [[antibacterial]], [[antifungal]], [[antiviral]], [[antiparasitic]], [[bradykinin]]-potentiating, and [[immunosuppressive]] drugs. As of 2020, no scorpion toxin-based drug is for sale, though [[chlorotoxin]] is being trialled for use against [[glioma]], a brain cancer.<ref name="Ahmadi Knerr Argemi Bordon p=118">{{cite journal |last1=Ahmadi |first1=Shirin |last2=Knerr |first2=Julius M. |last3=Argemi |first3=Lídia |last4=Bordon |first4=Karla C. F. |last5=Pucca |first5=Manuela B. |last6=Cerni |first6=Felipe A. |last7=Arantes |first7=Eliane C. |last8=Çalışkan |first8=Figen |last9=Laustsen |first9=Andreas H. |title=Scorpion Venom: Detriments and Benefits |journal=Biomedicines |publisher=MDPI AG |volume=8 |issue=5 |date=12 May 2020 |issn=2227-9059 |doi=10.3390/biomedicines8050118 |pmid=32408604 |pmc=7277529 |page=118|doi-access=free }}</ref>
 ===Consumption===
@@ Line 268: / Line 271: @@
 Scorpion and snake fighting Anglo-Saxon c 1050.jpg|"Scorpion and snake fighting", [[Cotton Vitellius|''Anglo-Saxon Herbal'']], c. 1050
 File:Sidney Hall - Urania's Mirror - Scorpio.jpg|The constellation [[Scorpius]], depicted in ''[[Urania's Mirror]]'' as "Scorpio", London, c. 1825
-File:Scorpion kilim motif.jpg|A [[Kilim motifs|scorpion motif]] (two types shown) was often woven into Turkish ''[[kilim]]'' flatweave carpets, for protection from their sting.<ref name="Erbek 1998"/>
+File:Scorpion kilim motif.jpg|A [[scorpion motif]] (two types shown) was often woven into Turkish ''[[kilim]]'' flatweave carpets, for protection from their sting.<ref name="Erbek 1998"/>
 </gallery>
 The scorpion is a culturally significant animal, appearing as a [[Motif (visual arts)|motif]] in art, especially in [[Islamic art]] in the Middle East.<ref name="Frembgen">{{cite journal |first=Jürgen Wasim |last=Frembgen |year=2004 |title=The Scorpion in Muslim Folklore |journal=[[Asian Folklore Studies]] |volume=63 |issue=1 |pages=95–123 |url=http://nirc.nanzan-u.ac.jp/nfile/631 |access-date=12 August 2014 |archive-date=16 March 2014 |archive-url=https://web.archive.org/web/20140316232038/http://nirc.nanzan-u.ac.jp/nfile/631 |url-status=live }}</ref>
-A [[Kilim motifs|scorpion motif]] is often woven into Turkish [[kilim]] flatweave carpets, for protection from their sting.<ref name="Erbek 1998">{{cite book |last=Erbek |first=Güran |title=Kilim Catalogue No. 1 |publisher=May Selçuk A. S. |edition=1st |date=1998 |at="Motifs", before page 1}}</ref> The scorpion is perceived both as an embodiment of evil and as a protective force such as a [[dervish]]'s powers to combat evil.<ref name="Frembgen"/> In Muslim folklore, the scorpion portrays [[human sexuality]].<ref name="Frembgen"/> Scorpions are used in [[folk medicine]] in South Asia, especially in [[antidote]]s for scorpion stings.<ref name="Frembgen"/>
+A [[scorpion motif]] is often woven into Turkish [[kilim]] flatweave carpets, for protection from their sting.<ref name="Erbek 1998">{{cite book |last=Erbek |first=Güran |title=Kilim Catalogue No. 1 |publisher=May Selçuk A. S. |edition=1st |date=1998 |at="Motifs", before page 1}}</ref> The scorpion is perceived both as an embodiment of evil and as a protective force such as a [[dervish]]'s powers to combat evil.<ref name="Frembgen"/> In Muslim folklore, the scorpion portrays [[human sexuality]].<ref name="Frembgen"/> Scorpions are used in [[folk medicine]] in South Asia, especially in [[antidote]]s for scorpion stings.<ref name="Frembgen"/>
-One of the earliest occurrences of the scorpion in culture is its inclusion, as ''[[Scorpio (astrology)|Scorpio]]'', in the 12 [[signs of the Zodiac]] by [[Babylonian astronomy|Babylonian astronomers]] during the [[Neo-Babylonian|Chaldean period]]. This was then taken up by western [[astrology]]; in astronomy the corresponding constellation is named [[Scorpius]].{{sfn|Polis|1990|p=462}}
+One of the earliest occurrences of the scorpion in culture is its inclusion, as ''[[Scorpio (astrology)|Scorpio]]'', in the 12 [[signs of the Zodiac]] by [[Babylonian astronomers]] during the [[Neo-Babylonian|Chaldean period]]. This was then taken up by western [[astrology]]; in astronomy the corresponding constellation is named [[Scorpius]].{{sfn|Polis|1990|p=462}}
 In [[ancient Egypt]], the goddess [[Serket]], who protected the [[Pharaoh]], was often depicted as a scorpion.<ref>{{cite web |url=https://www.britishmuseum.org/collection/term/BIOG55814 |title=Serqet |publisher=[[British Museum]] |access-date=10 December 2020 |archive-date=1 December 2020 |archive-url=https://web.archive.org/web/20201201174344/https://www.britishmuseum.org/collection/term/BIOG55814 |url-status=live }}</ref>
 In [[ancient Greece]], a warrior's shield sometimes carried a scorpion device, as seen in [[red-figure pottery]] from the 5th century BC.<ref>{{cite web |last1=attrib. to "The Achilles Painter" |title=Pelike |url=https://www.britishmuseum.org/collection/object/G_1856-1226-46 |publisher=[[British Museum]] |ref=1856,1226.46 |quote=in his left [hand] he carries a long spear and a shield with device in black silhouette of a scorpion to left. ... Made in: Attica, Greece ... Findspot: Nola, Italy |access-date=24 August 2020 |archive-date=3 October 2020 |archive-url=https://web.archive.org/web/20201003103339/https://www.britishmuseum.org/collection/object/G_1856-1226-46 |url-status=live }}</ref> In [[Greek mythology]], [[Artemis]] or [[Gaia]] sent a giant scorpion to kill the hunter [[Orion (mythology)|Orion]], who had said he would kill all the world's animals. Orion and the scorpion both became constellations; as enemies they were placed on opposite sides of the world, so when one rises in the sky, the other sets.<ref name="Kerényi 1974">{{cite book |last1=Kerényi |first1=C. |author1-link=Károly Kerényi |title=The Gods of the Greens |date=1974 |publisher=Thames and Hudson |isbn=978-0-500-27048-6 |page=203 |chapter=Stories of Orion}}</ref>{{sfn|Stockmann|Ythier|2010|p=179}} Scorpions are mentioned in the [[Bible]] and the [[Talmud]] as symbols of danger and maliciousness.{{sfn|Stockmann|Ythier|2010|p=179}}
-The [[Animal tale|fable]] of ''[[The Scorpion and the Frog]]'' has been interpreted as showing that vicious people cannot resist hurting others, even when it is not in their interests.<ref>{{cite journal |author=Takeda, A. |url=https://publikationen.uni-tuebingen.de/xmlui/bitstream/handle/10900/47008/pdf/Arata_Takeda_Blumenreiche_Handelswege.pdf |title=Blumenreiche Handelswege: Ost-westliche Streifzüge auf den Spuren der Fabel ''Der Skorpion und der Frosch'' |trans-title=Flowery Trade Routes: East-Western forays into the footsteps of the fable ''The Scorpion and the Frog'' |journal=Deutsche Vierteljahrsschrift für Literaturwissenschaft und Geistesgeschichte |language=de |year=2011 |volume=85 |issue=1 |pages=124–152 |doi=10.1007/BF03374756 |s2cid=170169337 |quotation=(German) Die Moral der Fabel besagt: Manche Menschen handeln von Natur aus mörderisch und selbst-mörderisch zugleich. (English) The moral of the fable says: Some people act naturally murderous and self-murderous at the same time. |access-date=9 December 2020 |archive-date=5 November 2020 |archive-url=https://web.archive.org/web/20201105152526/https://publikationen.uni-tuebingen.de/xmlui/bitstream/handle/10900/47008/pdf/Arata_Takeda_Blumenreiche_Handelswege.pdf |url-status=live }}</ref> More recently, the action in [[John Steinbeck]]'s 1947 novella ''[[The Pearl (novel)|The Pearl]]'' centers on a poor pearl fisherman's attempts to save his infant son from a scorpion sting, only to lose him to human violence.<ref>{{cite journal |last1=Meyer |first1=Michael |title=Diamond in the Rough: Steinbeck's Multifaceted Pearl |journal=The Steinbeck Review |date=2005 |volume=2 |issue=2 (Fall 2005) |pages=42–56 |jstor=41581982}}</ref> Scorpions have equally appeared in western artforms including film and poetry: the [[Surrealism|surrealist]] [[filmmaker]] [[Luis Buñuel]] made symbolic use of scorpions in his 1930 classic ''[[L'Age d'or]]'' (''The Golden Age''),<ref>{{cite book |first=Allen S. |last=Weiss |chapter=Between the sign of the scorpion and the sign of the cross: ''L'Age d'or'' |pages=[https://archive.org/details/isbn_9780262611213/page/159 159] |editor-first=Rudolf E. |editor-last=Kuenzli |year=1996 |title=Dada and Surrealist Film |publisher=[[MIT Press]] |isbn=978-0-262-61121-3 |chapter-url=https://books.google.com/books?id=ZDl6HsqFJwAC&pg=PA159 |url=https://archive.org/details/isbn_9780262611213/page/159 }}</ref> while [[Stevie Smith]]'s last collection of poems was entitled ''Scorpion and other Poems''.<ref>{{cite web |url=https://www.poetryfoundation.org/poets/stevie-smith |title=Stevie Smith: Bibliography |website=Poetry Foundation |access-date=1 July 2019 |archive-date=1 July 2019 |archive-url=https://web.archive.org/web/20190701135929/https://www.poetryfoundation.org/poets/stevie-smith |url-status=live }}</ref> A variety of [[martial arts]] films and video games have been entitled ''Scorpion King''.<ref>{{cite web |url=http://www.dvdtalk.com/reviews/11333/operation-scorpio/ |title=Operation Scorpio |last=Wallis |first=J. Doyle |work=[[DVD Talk]] |year=2004 |access-date=19 June 2015 |archive-date=20 June 2015 |archive-url=https://web.archive.org/web/20150620214728/http://www.dvdtalk.com/reviews/11333/operation-scorpio/ |url-status=live }}</ref><ref>{{cite web |title=The Scorpion King |website=[[Box Office Mojo]] |url=https://boxofficemojo.com/movies/?id=scorpionking.htm |access-date=4 January 2010 |archive-date=16 December 2009 |archive-url=https://web.archive.org/web/20091216212000/http://www.boxofficemojo.com/movies/?id=scorpionking.htm |url-status=live }}</ref><ref>{{cite web |last=Provo |first=Frank |url=http://www.gamespot.com/reviews/the-scorpion-king-sword-of-osiris-review/1900-2861730/ |title=The Scorpion King: Sword of Osiris Review |publisher=GameSpot |date=2002 |access-date=24 June 2020 |archive-date=29 June 2020 |archive-url=https://web.archive.org/web/20200629174813/https://www.gamespot.com/reviews/the-scorpion-king-sword-of-osiris-review/1900-2861730/ |url-status=live }}</ref>
+The [[Animal tale|fable]] of ''[[The Scorpion and the Frog]]'' has been interpreted as showing that vicious people cannot resist hurting others, even when it is not in their interests.<ref>{{cite journal |author=Takeda, A. |url=https://publikationen.uni-tuebingen.de/xmlui/bitstream/handle/10900/47008/pdf/Arata_Takeda_Blumenreiche_Handelswege.pdf |title=Blumenreiche Handelswege: Ost-westliche Streifzüge auf den Spuren der Fabel ''Der Skorpion und der Frosch'' |trans-title=Flowery Trade Routes: East-Western forays into the footsteps of the fable ''The Scorpion and the Frog'' |journal=Deutsche Vierteljahrsschrift für Literaturwissenschaft und Geistesgeschichte |language=de |year=2011 |volume=85 |issue=1 |pages=124–152 |doi=10.1007/BF03374756 |s2cid=170169337 |quotation=(German) Die Moral der Fabel besagt: Manche Menschen handeln von Natur aus mörderisch und selbst-mörderisch zugleich. (English) The moral of the fable says: Some people act naturally murderous and self-murderous at the same time. |access-date=9 December 2020 |archive-date=5 November 2020 |archive-url=https://web.archive.org/web/20201105152526/https://publikationen.uni-tuebingen.de/xmlui/bitstream/handle/10900/47008/pdf/Arata_Takeda_Blumenreiche_Handelswege.pdf |url-status=live }}</ref> More recently, the action in [[John Steinbeck]]'s 1947 novella ''[[The Pearl (novel)|The Pearl]]'' centers on a poor pearl fisherman's attempts to save his infant son from a scorpion sting, only to lose him to human violence.<ref>{{cite journal |last1=Meyer |first1=Michael |title=Diamond in the Rough: Steinbeck's Multifaceted Pearl |journal=The Steinbeck Review |date=2005 |volume=2 |issue=2 (Fall 2005) |pages=42–56 |jstor=41581982}}</ref> Scorpions have equally appeared in western artforms including film and poetry: the [[surrealist]] [[filmmaker]] [[Luis Buñuel]] made symbolic use of scorpions in his 1930 classic ''[[L'Age d'or]]'' (''The Golden Age''),<ref>{{cite book |first=Allen S. |last=Weiss |chapter=Between the sign of the scorpion and the sign of the cross: ''L'Age d'or'' |pages=[https://archive.org/details/isbn_9780262611213/page/159 159] |editor-first=Rudolf E. |editor-last=Kuenzli |year=1996 |title=Dada and Surrealist Film |publisher=[[MIT Press]] |isbn=978-0-262-61121-3 |chapter-url=https://books.google.com/books?id=ZDl6HsqFJwAC&pg=PA159 |url=https://archive.org/details/isbn_9780262611213/page/159 }}</ref> while [[Stevie Smith]]'s last collection of poems was entitled ''Scorpion and other Poems''.<ref>{{cite web |url=https://www.poetryfoundation.org/poets/stevie-smith |title=Stevie Smith: Bibliography |website=Poetry Foundation |access-date=1 July 2019 |archive-date=1 July 2019 |archive-url=https://web.archive.org/web/20190701135929/https://www.poetryfoundation.org/poets/stevie-smith |url-status=live }}</ref> A variety of [[martial arts]] films and video games have been entitled ''Scorpion King''.<ref>{{cite web |url=http://www.dvdtalk.com/reviews/11333/operation-scorpio/ |title=Operation Scorpio |last=Wallis |first=J. Doyle |work=[[DVD Talk]] |year=2004 |access-date=19 June 2015 |archive-date=20 June 2015 |archive-url=https://web.archive.org/web/20150620214728/http://www.dvdtalk.com/reviews/11333/operation-scorpio/ |url-status=live }}</ref><ref>{{cite web |title=The Scorpion King |website=[[Box Office Mojo]] |url=https://boxofficemojo.com/movies/?id=scorpionking.htm |access-date=4 January 2010 |archive-date=16 December 2009 |archive-url=https://web.archive.org/web/20091216212000/http://www.boxofficemojo.com/movies/?id=scorpionking.htm |url-status=live }}</ref><ref>{{cite web |last=Provo |first=Frank |url=http://www.gamespot.com/reviews/the-scorpion-king-sword-of-osiris-review/1900-2861730/ |title=The Scorpion King: Sword of Osiris Review |publisher=GameSpot |date=2002 |access-date=24 June 2020 |archive-date=29 June 2020 |archive-url=https://web.archive.org/web/20200629174813/https://www.gamespot.com/reviews/the-scorpion-king-sword-of-osiris-review/1900-2861730/ |url-status=live }}</ref>
-[[File:Mr-yoga-stretched-out-scorpion.jpg.png|thumb|[[Vrischikasana|Scorpion pose]] in [[yoga as exercise|yoga]] has one or both legs pointing forward over the head, like a scorpion's tail.<ref name="Iyengar 1991"/>]]
+[[File:Mr-yoga-stretched-out-scorpion.jpg.png|thumb|[[Scorpion pose]] in [[yoga as exercise|yoga]] has one or both legs pointing forward over the head, like a scorpion's tail.<ref name="Iyengar 1991"/>]]
-Since [[Classical era|classical times]], the scorpion with its powerful stinger has been used to provide a name for weapons. In the [[Roman army]], the [[Scorpio (weapon)|scorpio]] was a torsion siege engine used to shoot a projectile.<ref name="Vitruvius">[[Vitruvius]], ''De Architectura'', X:10:1–6.</ref> The British Army's [[FV101 Scorpion]] was an armored reconnaissance vehicle or light [[tank]] in service from 1972 to 1994.<ref name="Jane's">{{cite web |publisher=[[Jane's Information Group]] |title=Scorpion |url=http://www.janes.com/defence/land_forces/supplement/lav/lav_scorpion.shtml |access-date=16 November 2020 |archive-url=https://web.archive.org/web/20080221171530/https://www.janes.com/defence/land_forces/supplement/lav/lav_scorpion.shtml |archive-date=21 February 2008 |url-status=dead}}</ref> A version of the [[Matilda II]] tank, fitted with a [[mine flail|flail]] to clear [[Land mine|mines]], was named the Matilda Scorpion.<ref name="Fletcher2017">{{cite book |last=Fletcher |first=David |title=British Battle Tanks: British-made Tanks of World War II |url=https://books.google.com/books?id=H7EvDwAAQBAJ&pg=PA37 |year=2017 |publisher=Bloomsbury |isbn=978-1-4728-2003-7 |page=37 |access-date=27 June 2020 |archive-date=31 July 2020 |archive-url=https://web.archive.org/web/20200731185805/https://books.google.com/books?id=H7EvDwAAQBAJ&pg=PA37 |url-status=live }}</ref>
+Since [[classical times]], the scorpion with its powerful stinger has been used to provide a name for weapons. In the [[Roman army]], the [[Scorpio (weapon)|scorpio]] was a torsion siege engine used to shoot a projectile.<ref name="Vitruvius">[[Vitruvius]], ''De Architectura'', X:10:1–6.</ref> The British Army's [[FV101 Scorpion]] was an armored reconnaissance vehicle or light [[tank]] in service from 1972 to 1994.<ref name="Jane's">{{cite web |publisher=[[Jane's Information Group]] |title=Scorpion |url=http://www.janes.com/defence/land_forces/supplement/lav/lav_scorpion.shtml |access-date=16 November 2020 |archive-url=https://web.archive.org/web/20080221171530/https://www.janes.com/defence/land_forces/supplement/lav/lav_scorpion.shtml |archive-date=21 February 2008 |url-status=dead}}</ref> A version of the [[Matilda II]] tank, fitted with a [[mine flail|flail]] to clear [[Land mine|mines]], was named the Matilda Scorpion.<ref name="Fletcher2017">{{cite book |last=Fletcher |first=David |title=British Battle Tanks: British-made Tanks of World War II |url=https://books.google.com/books?id=H7EvDwAAQBAJ&pg=PA37 |year=2017 |publisher=Bloomsbury |isbn=978-1-4728-2003-7 |page=37 |access-date=27 June 2020 |archive-date=31 July 2020 |archive-url=https://web.archive.org/web/20200731185805/https://books.google.com/books?id=H7EvDwAAQBAJ&pg=PA37 |url-status=live }}</ref>
 Several<!--At least ten--> ships [[HMS Scorpion|of the Royal Navy]] and [[USS Scorpion|of the US Navy]] have been named ''Scorpion'' including [[HMS Scorpion (1803)|an 18-gun sloop]] in 1803,<ref>{{cite book |last=Winfield |first=Rif |title=British Warships in the Age of Sail 1793–1817: Design, Construction, Careers and Fates |publisher=Seaforth |year=2008 |isbn=978-1-86176-246-7 |page=291}}</ref> [[HMS Scorpion (1863)|a turret ship]] in 1863,<ref>{{cite book |last=Parkes |first=Oscar |title=British Battleships |publisher=Naval Institute Press |year=1990 |edition=reprint of the 1957 |isbn=1-55750-075-4 |pages=78–79}}</ref> [[USS Scorpion (PY-3)|a patrol yacht]] in 1898,<ref>{{cite journal |date=10 September 1896 |title=[untitled] |journal=Marine Review |volume=14 |issue=11 |location=Cleveland, Ohio |page=1}}</ref> [[HMS Scorpion (1910)|a destroyer]] in 1910,<ref>{{cite news |title=[untitled] |department=Naval and Military Intelligence |work=[[The Times]] |location=London |date=31 August 1910 |page=5 |url=https://www.newspapers.com/clip/64444910/scorpion-tbd/ |access-date=2 December 2020 |archive-date=31 May 2021 |archive-url=https://web.archive.org/web/20210531031226/https://www.newspapers.com/clip/64444910/scorpion-tbd/ |url-status=live }}</ref> and [[USS Scorpion (SSN-589)|a nuclear submarine]] in 1960.<ref>{{cite web |url=http://www.csp.navy.mil/othboats/589.htm |title=USS Scorpion (SSN 589) May 27, 1968&nbsp;– 99 Men Lost |date=2007 |website=[[United States Navy]] |access-date=9 April 2008 |url-status=dead |archive-url=https://web.archive.org/web/20080312204810/http://www.csp.navy.mil/othboats/589.htm |archive-date=12 March 2008 }}</ref>
 The scorpion has served as the name or symbol of products and brands including Italy's [[Abarth]] racing cars<ref>{{cite web |title=The History of Abarth's Logo |url=http://www.museodelmarchioitaliano.com/route1/abarth.php |publisher=Museo del marchio italiano |access-date=16 November 2020 |quote=The company logo has been the scorpion since the very start; Carlo Abarth did not only want it as a reference to his zodiac sign, but also because it was an original and hard to imitate logo. At the beginning the scorpion was free from any contour and featured the typo "Abarth & Co.- Torino". In 1954 a shield was added, as symbol of victory and passion; |archive-date=3 July 2020 |archive-url=https://web.archive.org/web/20200703181150/http://www.museodelmarchioitaliano.com/route1/abarth.php |url-status=live }}</ref> and a [[Montesa Honda|Montesa]] scrambler [[motorcycle]].<ref>{{cite web |last=Salvadori |first=Clement |title=Retrospective: 1974–1977 Montesa Cota 247-T |url=https://ridermagazine.com/2019/01/17/retrospective-1974-1977-montesa-cota-247-t/ |work=Rider Magazine |access-date=25 June 2020 |date=17 January 2019 |quote=Permanyer persisted, built larger engines, and in 1965 showed the 247cc engine (21 horsepower at 7,000 rpm) in a Scorpion motocrosser. |archive-date=27 June 2020 |archive-url=https://web.archive.org/web/20200627063031/https://ridermagazine.com/2019/01/17/retrospective-1974-1977-montesa-cota-247-t/ |url-status=live }}</ref>
-A hand- or forearm-balancing [[asana]] in modern [[yoga as exercise]] with the back arched and one or both legs pointing forward over the head in the manner of the scorpion's tail is called [[Vrischikasana|Scorpion pose]].<ref name="Budig 2012">{{cite journal |author=Anon |last2=Budig |first2=Kathryn |title=Kathryn Budig Challenge Pose: Scorpion in Forearm Balance |journal=[[Yoga Journal]] |date=1 October 2012 |url=https://www.yogajournal.com/practice/vrischika-in-pincha-mayurasana |access-date=25 June 2020 |archive-date=27 June 2020 |archive-url=https://web.archive.org/web/20200627182355/https://www.yogajournal.com/practice/vrischika-in-pincha-mayurasana |url-status=live }}</ref><ref name="Iyengar 1991">{{cite book |last=Iyengar |first=B. K. S. |author-link=B. K. S. Iyengar |title=[[Light on Yoga]] |publisher=Thorsons |year=1991 |isbn=978-0-00-714516-4 |oclc=51315708 |pages=386–388}}</ref>
+A hand- or forearm-balancing [[asana]] in modern [[yoga as exercise]] with the back arched and one or both legs pointing forward over the head in the manner of the scorpion's tail is called [[Scorpion pose]].<ref name="Budig 2012">{{cite journal |author=Anon |last2=Budig |first2=Kathryn |title=Kathryn Budig Challenge Pose: Scorpion in Forearm Balance |journal=[[Yoga Journal]] |date=1 October 2012 |url=https://www.yogajournal.com/practice/vrischika-in-pincha-mayurasana |access-date=25 June 2020 |archive-date=27 June 2020 |archive-url=https://web.archive.org/web/20200627182355/https://www.yogajournal.com/practice/vrischika-in-pincha-mayurasana |url-status=live }}</ref><ref name="Iyengar 1991">{{cite book |last=Iyengar |first=B. K. S. |author-link=B. K. S. Iyengar |title=[[Light on Yoga]] |publisher=Thorsons |year=1991 |isbn=978-0-00-714516-4 |oclc=51315708 |pages=386–388}}</ref>
 ==Notes==