Scorpion

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Scorpions
Temporal range: 435–0 
Ma
Early Silurian – present
Hottentotta tamulus from Mangaon, Maharashtra, India
Scientific classification Edit this 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

Scorpions are

genomic
studies.

Scorpions primarily prey on

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.

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

Allopalaeophonus, formerly called Palaeophonus hunteri, from the Silurian of Scotland[4]

Fossil record

Palaeophonus nuncius, a Silurian fossil from Sweden

Scorpion

Chactoidea and Buthoidea respectively, indicating that the crown group of modern scorpions had emerged by this time.[13]

Phylogeny

The Scorpiones are a

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

(

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

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

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
Heterometrus laoticus, the Vietnam forest scorpion, a member of the family Scorpionidae

Geographical distribution

Scorpions are found on all continents except

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

endemic to the littoral zone of rivers in Romania, occupy specialized niches.[38][39]

Morphology

Chelae;
8 = Moveable claw or Tarsus;
9 = Fixed claw or Manus;
10 = Stinger or Aculeus;
11 = Telson
(anus in previous joint);
12 = Opening of book lungs

Scorpions range in size from the 8.5 mm (0.33 in)

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

ganglia which may be a primitive trait.[48]

The pedipalp is a segmented, clawed

setae.[50] Depending on the species, the legs may have spines and spurs.[51]

Mesosoma

Ventral view: the pectines have a comblike structure in an inverted V shape.

The mesosoma or preabdomen is the broad part of the opisthosoma.

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

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

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

Stinger of an Arizona bark scorpion

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

Centruroides limpidus in its rocky shelter

Most scorpion species are

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.

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

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 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 128 cm/s (50 in/s) in a defensive strike.[75]

Mortality and defense

Scorpions may be attacked by other arthropods like ants, spiders,

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

solifugid

Scorpions generally prey on insects, particularly

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

metabolic rate, and a relatively inactive lifestyle. This enables some to survive six to twelve months of starvation.[87]

Mating

Male and female scorpion during promenade à deux

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

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

Scorpions glow a vibrant blue-green when exposed to certain wavelengths of

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

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

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

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

Pandinus imperator.[121]

Culture

  • Late period bronze figure of Isis-Serket
    Late period bronze figure of Isis-Serket
  • "Scorpion and snake fighting", Anglo-Saxon Herbal, c. 1050
    "Scorpion and snake fighting",
    Anglo-Saxon Herbal
    , c. 1050
  • The constellation Scorpius, depicted in Urania's Mirror as "Scorpio", London, c. 1825
    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]
    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

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

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

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

Scorpion pose.[147][136]

Notes

  1. 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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