scorpion

Scorpion Temporal range: Silurian–Recent PreЄ Є O S D C P T J K Pg N
Asian forest scorpion in Khao Yai National Park, Thailand
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Chelicerata
Class:	Arachnida
Subclass:	Dromopoda
Order:	Scorpiones C. L. Koch, 1837
Superfamilies
Buthoidea Chaeriloidea Chactoidea Iuroidea Pseudochactoidea Scorpionoidea See classification for families.

Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger. Scorpions range in size from 9 mm (Typhlochactas mitchelli) to 21 cm (Hadogenes troglodytes).^[1]

Scorpions are found widely distributed over all continents, except Antarctica, in a variety of terrestrial habitats except the high latitude tundra. Scorpions number about 1,752 described species,^[2] with 13 extant families recognised to date. The taxonomy has undergone changes and is likely to change further, as a number of genetic studies are bringing forth new information.

Though the scorpion has a fearsome reputation as venomous, only about 25 species are known to have venom capable of killing a human being.^[3]

Etymology

The word scorpion is thought to have originated in Middle English between 1175 and 1225 AD from Old French skorpiō,^[4] or from Italian scorpione, both derived from the Latin word scorpio,^[5] which in turn has its roots in the Greek word σκορπίος – skorpíos.^[6]

Geographical distribution

Scorpions are found on all major land masses except Antarctica. Scorpions did not occur naturally in Great Britain, New Zealand and some of the islands in Oceania, but have now been accidentally introduced in some of these places by human trade and commerce.^[3]:249 The greatest diversity of scorpions in the Northern Hemisphere is to be found in the subtropical areas lying between latitudes 23° N and 38° N. Above these latitudes, the diversity decreases, with the northernmost occurrence of scorpions being the northern scorpion Paruroctonus boreus at 50° N.^[3]:251

Today, scorpions are found in virtually every terrestrial habitat, including high elevation mountain-tops, caves and intertidal zones, with the exception of boreal ecosystems such as the tundra, high-altitude taiga and the permanently snow-clad tops of some mountains.^{[3]:251–252[7]} As regards microhabitats, scorpions may be ground-dwelling, tree-living, lithophilic (rock-loving) or psammophilic (sand-loving); some species such as Vaejovis janssi are versatile and found in every type of habitat in Baja California while others occupy specialised niches such as Euscorpius carpathicus which occupies the littoral zone of the shore.^[8]

Five colonies of scorpions (Euscorpius flavicaudis) have established themselves in Sheerness on the Isle of Sheppey in the United Kingdom.^[9] This small population has been resident since the 1860s, having probably arrived with imported fruit from Africa. This scorpion species is small and completely harmless to humans. This marks the northernmost limit in the world where scorpions live in the wild.^[10][11]

Classification

There are thirteen families and about 1,400 described species and subspecies of scorpions. In addition, there are 111 described taxa of extinct scorpions.^[12]

This classification is based on that of Soleglad & Fet (2003),^[13] which replaced the older, unpublished classification of Stockwell.^[14] Additional taxonomic changes are from papers by Soleglad et al. (2005).^[15][16]

Systematics

The following classification covers extant taxa to the rank of family.

Order Scorpiones

• • Infraorder Orthosterni Pocock, 1911
    • Parvorder Pseudochactida Soleglad et Fet, 2003
      • Superfamily Pseudochactoidea Gromov, 1998
        • Family Pseudochactidae Gromov, 1998
    • Parvorder Buthida Soleglad et Fet, 2003
      • Superfamily Buthoidea C. L. Koch, 1837
        • Family Buthidae C. L. Koch, 1837 (thick-tailed scorpions)
        • Family Microcharmidae Lourenço, 1996
    • Parvorder Chaerilida Soleglad et Fet, 2003
      • Superfamily Chaeriloidea Pocock, 1893
        • Family Chaerilidae Pocock, 1893
    • Parvorder Iurida Soleglad et Fet, 2003
      • Superfamily Chactoidea Pocock, 1893
        • Family Chactidae Pocock, 1893
        • Family Euscorpiidae Laurie, 1896
        • Family Superstitioniidae Stahnke, 1940
        • Family Vaejovidae Thorell, 1876
      • Superfamily Iuroidea Thorell, 1876
        • Family Caraboctonidae Kraepelin, 1905 (hairy scorpions)
        • Family Iuridae Thorell, 1876
      • Superfamily Scorpionoidea Latreille, 1802
        • Family Bothriuridae Simon, 1880
        • Family Hemiscorpiidae Pocock, 1893 (= Ischnuridae, =Liochelidae) (rock scorpions, creeping scorpions, or tree scorpions)
        • Family Scorpionidae Latreille, 1802 (burrowing scorpions or pale-legged scorpions)

Fossil record

Scorpions have been found in many fossil records, including marine Silurian deposits, coal deposits from the Carboniferous Period and in amber. The oldest known scorpions lived around 430 million years ago in the Silurian period, on the bottom of shallow tropical seas.^[17] These first scorpions had gills instead of the present forms' book lungs. Currently, 111 fossil species of scorpion are known.^[12] Unusually for arachnids, there are more species of Palaeozoic scorpion than Mesozoic or Cenozoic ones.

The eurypterids, marine creatures which lived during the Paleozoic era, share several physical traits with scorpions and may be closely related to them. Various species of Eurypterida could grow to be anywhere from 10 centimetres (3.9 in) to 2.5 metres (8.2 ft) in length.^[18] However, they exhibit anatomical differences marking them off as a group distinct from their Carboniferous and Recent relatives. Despite this, they are commonly referred to as "sea scorpions".^[19] Their legs are thought to have been short, thick, tapering and to have ended in a single strong claw; it appears that they were well-adapted for maintaining a secure hold upon rocks or seaweed against the wash of waves, like the legs of a shore crab.

Anatomy

Scorpion anatomy:
1 = Cephalothorax or Prosoma;
2 = Abdomen or Mesosoma;
3 = Tail or Metasoma;
4 = Claws or Pedipalps
5 = Legs;
6 = Mouth parts or Chelicerae;
7 = pincers or Chelae;
8 = Moveable claw or Tarsus;
9 = Fixed claw or Manus;
10 = Sting or Telson;
11 = Anus.

Barb of an Arizona bark scorpion

The body of a scorpion is divided into three parts (tagmata): the head (cephalothorax), the abdomen (mesosoma) and the tail (metasoma).^[3]:10

Cephalothorax

The cephalothorax, also called the prosoma, comprises the carapace, eyes, chelicerae (mouth parts), pedipalps (the pedipalps of scorpions have chelae commonly called claws or pincers) and four pairs of walking legs. The scorpion's exoskeleton is thick and durable, providing good protection from predators. 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. The position of the eyes on the cephalothorax depends in part on the hardness or softness of the soil upon which they spend their lives.^[20] ` The pedipalp is a segmented, chelate (clawed) appendage used for prey immobilization, defense, and sensory purposes. The segments of the pedipalp (from closest to the body outwards) are coxa, trochanter, femur (humerus), patella, tibia (including the fixed claw and the manus) and tarsus (moveable claw).^[21] A scorpion has darkened or granular raised linear ridges, called "keels" or carinae on the pedipalp segments and on other parts of the body, which are useful taxonomically.^[3]:12

Mesosoma

The abdomen, also called the opisthosoma, consists of seven segments (somites), each covered dorsally by a sclerotosed plate (tergum) and also ventrally for segments 3 to 7. The first abdominal segment bears a pair of genital opercula which cover the gonopore. Segment 2 consists of the basal plate with the pectines. Each of the mesosomal segments 3 to 7 have a pair of spiracles which are the openings for the scorpion's respiratory organs, known as book lungs. The spiracle openings may be slits, circular, elliptical, or oval.^[3]:13–15

Metasoma

The metasoma, the scorpion's tail, comprises five caudal segments (the first tail segment looks like a last mesosoman segment), and sixth bearing the telson (the sting). The telson, in turn, consists of the vesicle, which holds a pair of venom glands, and the hypodermic aculeus, the venom-injecting barb.

On rare occasions, scorpions can be born with two metasomata (tails). Two-tailed scorpions are not a different species, merely a genetic abnormality.^[22]

Fluorescence

A scorpion under a black light. In normal lighting, this scorpion appears black.

Scorpions are also known to glow when exposed to certain wavelengths of ultraviolet light such as that produced by a black light, due to the presence of fluorescent chemicals in the cuticle. One fluorescent component is now known to be beta-carboline.^[23] A hand-held UV 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.^[23] This fluorescence may have an active role in scorpion light detection.^[24]

Biology

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Scorpions have quite variable lifespans and the actual lifespan of most species is not known. The age range appears to be approximately 4–25 years (25 years being the maximum reported life span in the species Hadrurus arizonensis). Lifespan of Hadogenes species in the wild is estimated at 25–30 years.

Scorpions prefer to live in areas where the temperatures range from 20 to 37 °C (68 to 99 °F), but may survive freezing temperatures to the desert heat.^[25][26] Scorpions of the genus Scorpiops living in high Asian mountains, bothriurid scorpions from Patagonia and small Euscorpius scorpions from Central Europe can all survive winter temperatures of about −25 °C (−13 °F). In Repetek (Turkmenistan), there live seven species of scorpions (of which Pectinibuthus birulai is endemic) in temperatures which vary from -31–50 °C (-24–122 °F).^[27]

They are nocturnal and fossorial, finding shelter during the day in the relative cool of underground holes or undersides of rocks and coming out at night to hunt and feed. Scorpions exhibit photophobic behavior, primarily to evade detection by their predators such as birds, centipedes, lizards, mice, possums, and rats.^[28]

Scorpions are opportunistic predators of small arthropods, although the larger kinds have been known to kill small lizards and mice. The large pincers are studded with highly sensitive tactile hairs, and the moment an insect touches these, they use their chelae (pincers) to catch the prey. Depending on the toxicity of their venom and size of their claws, they will then either crush the prey or inject it with neurotoxic venom. This will kill or paralyze the prey so the scorpion can eat it. Scorpions have a relatively unique style of eating using chelicerae, small claw-like structures that protrude from the mouth that are unique to the Chelicerata among arthropods. The chelicerae, which are very sharp, are used to pull small amounts of food off the prey item for digestion into a pre-oral cavity below the chelicerae and carapace. Scorpions can only ingest food in a liquid form; they have external digestion. The digestive juices from the gut are egested onto the food and the digested food sucked in liquid form. Any solid indigestible matter (fur, exoskeleton, etc.) is trapped by setae in the pre-oral cavity, which is ejected by the scorpion.^{[3]:296–297}

Scorpions can consume huge amounts of food at one sitting. They have a very efficient food storage organ and a very low metabolic rate combined with a relatively inactive lifestyle. This enables scorpions to survive long periods when deprived of food; some are able to survive 6 to 12 months of starvation.^{[3]:297–298} Scorpions excrete very little; their waste consists mostly of insoluble nitrogenous waste such as xanthine, guanine and uric acid.^[8]

Reproduction

Most scorpions reproduce sexually, and most species have male and female individuals. However, some species, such as Hottentotta hottentotta, Hottentotta caboverdensis, Liocheles australasiae, Tityus columbianus, Tityus metuendus, Tityus serrulatus, Tityus stigmurus, Tityus trivittatus, and Tityus urugayensis, reproduce through parthenogenesis, a process in which unfertilised eggs develop into living embryos. Parthenogenic reproduction starts following the scorpion's final moult to maturity and continues thereafter.

Sexual reproduction is accomplished by the transfer of a spermatophore from the male to the female; scorpions possess a complex courtship and mating ritual to effect this transfer. Mating starts with the male and female locating and identifying each other using a mixture of pheromones and vibrational communication. Once they have satisfied the other that they are of opposite sex and of the correct species, mating can commence.

The courtship starts with the male grasping the female's pedipalps with his own; the pair then perform a "dance" called the "promenade à deux". In this "dance," the male leads the female around searching for a suitable place to deposit his spermatophore. The courtship ritual can involve several other behaviours such as juddering and a cheliceral kiss, in which the male's chelicerae – pincers – grasp the female's in a smaller more intimate version of the male's grasping the female's pedipalps and in some cases injecting a small amount of his venom into her pedipalp or on the edge of her cephalothorax,^[29] probably as a means of pacifying the female.

When the male has identified a suitable location, he deposits the spermatophore and then guides the female over it. This allows the spermatophore to enter her genital opercula, which triggers release of the sperm, thus fertilizing the female. The mating process can take from 1 to 25+ hours and depends on the ability of the male to find a suitable place to deposit his spermatophore. If mating goes on for too long, the female may eventually lose interest, breaking off the process.

Once the mating is complete, the male and female will separate. The male will generally retreat quickly, most likely to avoid being cannibalised by the female, although sexual cannibalism is infrequent with scorpions.

Birth and development

Compsobuthus werneri female with young

Unlike the majority of species in the class arachnid, which are oviparous, scorpions seem to be universally viviparous. The young are born one by one after hatching and expelling the embryonic membrane, if any, and the brood is carried about on its mother's back until the young have undergone at least one molt. Before the first molt, scorplings cannot survive naturally without the mother, since they depend on her for protection and to regulate their moisture levels. Especially in species which display more advanced sociability (e.g. Pandinus spp.), the young/mother association can continue for an extended period of time. The size of the litter depends on the species and environmental factors, and can range from two to over a hundred scorplings. The average litter however, consists of around 8 scorplings.^[30]

The young generally resemble their parents. Growth is accomplished by periodic shedding of the exoskeleton (ecdysis). A scorpion's developmental progress is measured in instars (how many moults it has undergone). Scorpions typically require between five and seven moults to reach maturity. Moulting commences with a split in the old exoskeleton just below the edge of the carapace (at the front of the prosoma). The scorpion then emerges from this split; the pedipalps and legs are first removed from the old exoskeleton, followed eventually by the metasoma. When it emerges, the scorpion's new exoskeleton is soft, making the scorpion highly vulnerable to attack. The scorpion must constantly stretch while the new exoskeleton hardens to ensure that it can move when the hardening is complete. The process of hardening is called sclerotization. The new exoskeleton does not fluoresce; as sclerotization occurs, the fluorescence gradually returns.

Relationship with humans

Scorpion sting and venom

All known scorpion species possess venom and use it primarily to kill or paralyze their prey so that it can be eaten; in general, it is fast-acting, allowing for effective prey capture. It is also used as a defense against predators. The venom is a mixture of compounds (neurotoxins, enzyme inhibitors, etc.) each not only causing a different effect, but possibly also targeting a specific animal. Each compound is made and stored in a pair of glandular sacs and is released in a quantity regulated by the scorpion itself. Of the 1000+ known species of scorpion, only 25 have venom that is dangerous to humans; most of those belong to the family Buthidae.^[8][31]

First aid

First aid for scorpion stings is generally symptomatic. It includes strong analgesia, either systemic (opiates or paracetamol) or locally applied (such as a cold compress). Hypertensive crises are treated with anxiolytics and vasodilators.^[32]

Medical use

The deathstalker has powerful venom.

The key ingredient of the venom is a scorpion toxin protein.

Short chain scorpion toxins constitute the largest group of potassium (K⁺) channel blocking peptides; an important physiological role of the KCNA3 channel, also known as K_V1.3, is to help maintain large electrical gradients for the sustained transport of ions such as Ca²⁺ that controls T lymphocyte (T cell) proliferation. Thus K_V1.3 blockers could be potential immunosuppressants for the treatment of autoimmune disorders (such as rheumatoid arthritis, inflammatory bowel disease and multiple sclerosis).^[33]

The venom of Uroplectes lineatus is clinically important in dermatology.^[34]

Toxins being investigated include:

• Chlorotoxin is a 36-amino acid peptide found in the venom of the deathstalker scorpion (Leiurus quinquestriatus) which blocks small-conductance chloride channels.^[35] The fact that chlorotoxin binds preferentially to glioma cells has allowed the development of new methods, that still are under investigation, for the treatment and diagnosis of several types of cancer.^[36]
• Maurotoxin from the venom of the Tunisian Scorpio maurus palmatus
• A number of antimicrobial peptides have also been found in the venom of Mesobuthus eupeus. Meucin-13 and Meucin-18 exhibited extensive cytolytic effects on bacteria, fungi and yeasts.^[37] Furthermore, Meucin-24 and Meucin-25, first identified from genetic sequences expressed in their venom gland, were shown to selectively kill Plasmodium falciparum and inhibit the development of Plasmodium berghei, both malaria parasites, but do not harm mammalian cells. These two venom-derived proteins are therefore attractive candidates for the development of anti-malarial drugs.^[38]

Iranian researchers have also reported that the venom of M. eupeus has anti-inflammatory properties and is effective as an anti-arthritis treatment in an experimental species model, but the mechanism of action is unknown.^[39]

Consumption

Eating scorpions in Beijing, China

Scorpion and snake wine

Fried scorpion is a traditional dish from Shandong, China.^[40]

As a part of Chinese medicine, scorpion wine and snake wine are used as analgesic and antidote.

In culture

• One of earliest occurrences of the scorpion in culture is its inclusion, as Scorpio, in the twelve signs of the series of constellations known as the Zodiac by Babylonian astronomers during the Chaldean period.^[3]:462

• In North Africa and South Asia, the scorpion is a significant animal culturally which appears as a motif in art, especially in Islamic art in the Middle East.^[41] It is perceived both as an embodiment of evil as well as a protective force which counters evil, such as a dervish's powers to combat evil.^[41] In another context, the scorpion portrays human sexuality.^[41] Scorpions are used in folk medicine in South Asia especially in antidotes for scorpion stings.^[41]

• In ancient Egypt the goddess Serket was often depicted as a scorpion, one of several goddesses who protected the Pharaoh.

• Surrealist filmmaker Luis Buñuel makes notable symbolic use of scorpions in his 1930 classic L'Age d'or (The Golden Age).^[42]

• The first two pages of Ian Fleming's novel Diamonds Are Forever are told from the point of view of an African scorpion which kills and eats a beetle and is then casually crushed and killed itself, by one of the villains whom James Bond would later confront and eventually crush.^[43]

See also

Arthropods portal

• Palaeophonus, a Devonian scorpion
• Paraisobuthus, a Carboniferous scorpion

References

External links

Wikispecies has information related to: Scorpiones

Wikimedia Commons has media related to: Scorpiones

• Biolumunescence in scorpions
• Scorpion anatomy
• Scorpions in Qatar
• Caring for a Pet Scorpion
• Holland, Ben (2008). "Scorpion Sweepers". http://www.scorpsweep.com/facts.html. 
• Bokma, John. "Scorpion detection using UV LEDs". http://johnbokma.com/pet/scorpion/detection-using-uv-leds.html. 
• "Scorpions". DesertUSA.com and Digital West Media, Inc.. http://www.desertusa.com/oct96/du_scorpion.html. 
• Rehak, Ondrej. "Photogallery - 36 species of scorpions". http://www.spidy.goliathus.com/english/gallery-scorpions.php. 
• Britt, Robert Roy (June 27, 2006). "Scorpion Venom Tested as Brain Cancer Treatment". LiveScience (Imaginova). http://www.livescience.com/health/060627_scorpion_venom.html. 
• Maney, Daniel (2006). "Beginner species of exotic pets". http://www.exotic-pets.eu/beginner.php. 
• "Scorpions - General information, care sheets, and more". TheInverts.com. http://www.theinverts.com/scorpioninformation.html. 
• "Scorpions. What is truth and what is wrong". http://knol.google.com/k/luis-f-de-armas/scorpions-what-is-truth-and-what-is/kbg2jg4ueep9/3#. 
• "Scorpions. The Antillean or West Indian fauna". http://knol.google.com/k/luis-f-de-armas/scorpions-the-antillean-or-west-indian/kbg2jg4ueep9/6#. 
• "Pet Scorpion Information and facts | Scorpion Picture Guide". http://www.scorpionpictureguide.com/. 
• Cuevas, Moises, MD (2009). "Scorpions in Sayulita". http://sayulitadoctor.com/2009/03/05/scorpions-in-sayulita/. 
• Scorpions from the Chihuahuan Desert Region of Mexico and the United States