Scorpion

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Scorpion
Asian forest scorpion (Heterometrus spinifer) 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
Pseudochactoidea Buthoidea Chaeriloidea Chactoidea Iuroidea Scorpionoidea See classification for families.

Scorpions are eight-legged carnivorous arthropods, members of the order Scorpiones within the class Arachnida. There are about 2000 species of scorpions, found widely distributed south of 49° N, except New Zealand and Antarctica. The northernmost part of the world where scorpions live in the wild is Sheerness on the Isle of Sheppey in the UK, where a small colony of Euscorpius flavicaudis has been resident since the 1860s.[1][2]

Contents 1 Anatomy 1.1 Cephalothorax 1.2 Mesosoma 1.3 Metasoma 2 Reproduction 3 Birth and development 4 Life and habits 5 Venom 6 Fossil record 7 Geographical distribution 8 Suicide misconception 9 Ultraviolet light 10 Classification 11 Cultural symbolism 12 See also 13 References 14 External links

[edit] Anatomy

The body of a scorpion is divided into two parts: the cephalothorax (also called the prosoma) and the abdomen (opisthosoma). The abdomen consists of the mesosoma and the metasoma.

[edit] Cephalothorax

The cephalothorax, also called the prosoma, is the scorpion's “head”, comprising the carapace, eyes, chelicerae (mouth parts), pedipalps (claws) and four pairs of walking legs. The scorpion's exoskeleton is thick and durable, providing good protection from predators.

[edit] Mesosoma

The mesosoma, the front half of the abdomen, is made up of six segments. The first segment contains the sexual organs as well as a pair of vestigial and modified appendages forming a structure called the genital operculum. The second segment bears a pair of featherlike sensory organs known as the pectines; the final four segments each contain a pair of book lungs. The mesosoma is armored with chitinous plates, known as tergites on the upper surface and sternites on the lower surface.

[edit] Metasoma

The metasoma, the scorpion's tail, comprises six segments (the first tail segment looks like a last mesosoman segment), the last containing the scorpion's anus and 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.^[1]

[edit] 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 unfertilized 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 each 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 reality this is the male leading 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--clawlike mouthparts--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,^[2] probably as a means of pacifying the female.

When he 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 cannibalized by the female, although sexual cannibalism is infrequent with scorpions.

[edit] Birth and development

Unlike the majority of arachnid species, scorpions are viviparous. The young are born one by one, and the brood is carried about on its mother's back until the young have undergone at least one moult. Before the first moult, 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.^[3]

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 is effected by means of a split in the old exoskeleton which takes place 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.

[edit] Life and habits

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°C to 37°C (68°F to 99°F), but may survive from freezing temperatures to the desert heat.^[4] Scorpions of the genus Scorpiops living in high Asian mountains, bothriurid scorpions from Patagonia and small Euscorpius scorpions from middle Europe can all survive winter temperatures of about -25°C.

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.^[5]

Scorpions are opportunistic predators of small arthropods and insects. They use their chelae (pincers) to catch the prey initially. 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. Scorpions can only digest food in a liquid form; any solid matter (fur, exoskeleton, etc) is disposed of by the scorpion.

[edit] Venom

All scorpion species possess venom. In general, scorpion venom is described as neurotoxic in nature. One exception to this generalization is Hemiscorpius lepturus which possesses cytotoxic venom. The neurotoxins consist of a variety of small proteins as well as sodium and potassium cations, which serve to interfere with neurotransmission in the victim. Scorpions use their venom to kill or paralyze their prey so that it can be eaten; in general it is fast-acting, allowing for effective prey capture. The effects of the sting can be severe.

Scorpion venoms are optimized for action upon other arthropods and therefore most scorpions are relatively harmless to humans; stings produce only local effects (such as pain, numbness or swelling). A few scorpion species, however, mostly in the family Buthidae, can be dangerous to humans. Among the most dangerous are Leiurus quinquestriatus, otherwise ominously known as the deathstalker, which has the most potent venom in the family, and members of the genera Parabuthus, Tityus, Centruroides, and especially Androctonus, which also have powerful venom. The scorpion which is responsible for the most human deaths is Androctonus australis, or the yellow fat-tailed scorpion, of North Africa. The toxicity of A. australis's venom is roughly half that of L. quinquestriatus, but despite a common misconception A. australis does not inject noticeably more venom into its prey. The higher death count is simply due to its being found more commonly, especially near humans. Human deaths normally occur in the young, elderly, or infirm; scorpions are generally unable to deliver enough venom to kill healthy adults. Some people, however, may be allergic to the venom of some species. Depending on the severity of the allergy, the scorpion's sting may cause anaphylaxis and death. A primary symptom of a scorpion sting is numbing at the injection site, sometimes lasting for several days. Scorpions are generally harmless and timid, and only voluntarily use their sting for killing prey, defending themselves or in territorial disputes with other scorpions. Generally, they will run from danger or remain still.

It should be noted that the family Buthidae, while containing perhaps the highest number of dangerous species, also contains many species that are not thought to be medically significant.

Scorpions are able to regulate how much venom is injected with each sting using striated muscles in the stinger, the usual amount being between 0.1 and 0.6 mg. There is also evidence to suggest that scorpions restrict the use of their venom using it only to subdue large prey, or prey that struggles. It has been found that scorpions have two types of venom: a translucent, weaker venom designed to stun only, and an opaque, more potent venom designed to kill heavier threats. This is likely because it is expensive in terms of energy for a scorpion to produce venom, and because it may take several days for a scorpion to replenish its venom supply once it has been exhausted.^[6][7]

There is currently no scorpion equivalent of the Schmidt Sting Pain Index, because nobody has yet classified the levels of pain inflicted by different scorpion stings. This is probably because of the risk involved with some species, such as Androctonus australis or Leiurus quinquestriatus. However, envenomation by a mildly venomous species like Pandinus imperator is similar to a bee sting in terms of the pain and swelling that results. A sting on the thumb from a relatively non-dangerous scorpion often feels like the victim has accidentally struck their thumb with a hammer whilst driving in a nail. A sting on the thumb from a truly dangerous scorpion can feel much worse, as though the victim had hammered a nail right through their thumb. It should be noted that the physical effects of a sting from a medically significant scorpion are not limited to the pain inflicted: there can be bradycardia, tachycardia or in severe cases pulmonary edema.

The stings of North American scorpions are rarely serious and usually result in pain, minimal swelling, tenderness, and warmth at the sting site. However, the Arizona bark scorpion Centruroides sculpturatus, which is found in Arizona and New Mexico and on the California side of the Colorado River, has a much more toxic sting. The sting is painful, sometimes causing numbness or tingling in the area around the sting. Serious symptoms are more common in children and include abnormal head, eye, and neck movements; increased saliva production; sweating; and restlessness. Some people develop severe involuntary twitching and jerking of muscles. Breathing difficulties may occur.

The stings of most North American scorpions require no special treatment. Placing an ice cube on the wound reduces pain, as does an ointment containing a combination of an antihistamine, an analgesic, and a corticosteroid. Centruroides stings that result in serious symptoms may require the use of sedatives, such as midazolam, given intravenously. Centruroides antivenom rapidly relieves symptoms, but it may cause a serious allergic reaction or serum sickness. The antivenom is available only in Arizona. In Trinidad the leaf juice of Eclipta prostrata is used for scorpion stings. Any effect of plants that are used against scorpion stings may be due to symptomatic relief – analgesic, anti-inflammatory, antipruritic effects, in addition to other biological activities. Some compounds from plants used for general inflammation also inhibit enzymes (like phospholipase A2) from snake and scorpion venom. Some of these plant compounds are hypolaetin-8-glucoside and related flavanoids.

Professor Moshe Gueron was the first to investigate the cardiovascular effects of a severe scorpion sting.^[8] Thousands of stung patients were reviewed. Thirty-four patients with severe scorpion stings were reviewed and pertinent data related to the cardiovascular system such as hypertension, peripheral vascular collapse, congestive heart failure or pulmonary edema were analyzed. The electrocardiograms of 28 patients were reviewed; 14 patients showed "early myocardial infarction-like" pattern. The urinary catecholamine metabolites were investigated in 12 patients with scorpion sting. Vanylmandelic acid was elevated in seven patients and the total free epinephrine and norepinephrine in eight. Six of these 12 patients displayed the electrocardiographic "myocardial infarction-like" pattern. Nine patients died and the pathologic lesions of the myocardium were reviewed in seven. Also, Gueron reported five cases of Severe Myocardial damage and heart failure in Scorpion sting from Beer-Sheba, Israel. He described hypertension, pulmonary oedema with hypertension, hypotension, pulmonary oedema with hypotension and rhythm disturbances as five different syndromes that may dominate the clinical picture in scorpion sting victim. He suggested that all patients with cardiac symptoms should be admitted to an intensive cardiac unit. A few years later, in 1990, he reported poor contractility with low ejection fraction, decreased systolic left ventricular performance, lowered fractional percentage shortening observed in echocardiographic and radionuclide angiographic study. Gueron was questioned regarding the value of giving antivenom, and he replied that although it is freely available, all cases of scorpion sting are treated without it, and there had not been a single fatality in 1989.^[9]

[edit] Fossil record

Scorpions have been found in many fossil records, including marine Silurian deposits, coal deposits from the Carboniferous Period and in amber. They are thought to have existed in some form since about 425–450 million years ago. They are believed to have an oceanic origin, with gills and a claw-like appendage that enabled them to hold onto rocky shores or seaweed, although the assumption that the oldest scorpions were aquatic has been questioned. Currently, 111 fossil species of scorpion are known. 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 cm (4 in) to 3 m (9.75 ft) in length. 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."^[10] 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 shore-crab.

[edit] Geographical distribution

Scorpions are almost universally distributed south of 49° N, and their geographical distribution shows in many particulars a close and interesting correspondence with that of the mammals, including their entire absence from New Zealand. The facts of their distribution are in keeping with the hypothesis that the order originated in the northern hemisphere and migrated southwards into the southern continent at various epochs, their absence from the countries to the north of the above-mentioned latitudes being due, no doubt, to the comparatively recent glaciation of those areas. When they reached Africa, Madagascar was part of that continent; but their arrival in Australia was subsequent to the separation of New Zealand from the Austro-Malayan area to the north of it.

In the United States, scorpions are most common in southern Arizona and in a swath of land extending through central Texas and central Oklahoma. The common striped scorpion, Centruroides vittatus, reaches from northwest Mexico to southern Colorado, Kansas, southern Missouri, and Mississippi and Louisiana. Species of the genus Vaejovis are found from Florida north to Maryland, the Carolinas, and Tennessee, and as far west as Oregon and California. Paruroctonus boreus is found through the Northwest U.S. and into Canada (Southern Saskatchewan, Southern Alberta and the Okanagan Valley of British Columbia). Scorpions can be found in 31 different states in the U.S., including Hawaii (Isometrus maculatus).

Five colonies of scorpions (Euscorpius flavicaudis) have established themselves in southern England having probably arrived with imported fruit from Africa, but the number of colonies could be lower now because of the destruction of their habitats. This scorpion species is small and completely harmless to humans.

[edit] Suicide misconception

The belief that scorpions commit suicide by stinging themselves to death when surrounded by fire (or when given alcohol) is of considerable antiquity and is often prevalent where these animals exist. It is nevertheless untrue since the venom has no effect on the scorpion itself, nor on any member of the same species (unless the venom is injected directly into the scorpion's nerve ganglion—quite an unlikely event outside of the laboratory). The misconception may derive from the fact that scorpions are poikilotherms (cold-blooded): when exposed to intense heat their metabolic processes malfunction. This causes the scorpion to spasm wildly and this spasming may appear as if the scorpion is stinging itself.

[edit] Ultraviolet light

Scorpions are also known to glow when exposed to certain wavelengths of ultraviolet light such as that produced by a blacklight, due to the presence of fluorescent chemicals in the cuticle. These have been stated to be quinones, but Stachel et al (1999) identified beta carboline as a principal fluorescent component. A hand-held UV lamp has long been a standard tool for nocturnal field surveys of these animals (eg Hadley & Williams 1968).

[edit] Classification

This classification is based on that of Soleglad & Fet (2003),^[11] which replaced the older, unpublished classification of Stockwell.^[12] Additional taxonomic changes are from Soleglad et al. (2005).^[13]

• 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
          • Subfamily Chactinae Pocock, 1893
            • Tribe Chactini Pocock, 1893
            • Tribe Nullibrotheini Soleglad et Fet, 2003
          • Subfamily Brotheinae Simon, 1879
            • Tribe Belisariini Lourenço, 1998
            • Tribe Brotheini Simon, 1879
              • Subtribe Brotheina Simon, 1879
              • Subtribe Neochactina Soleglad et Fet, 2003
          • Subfamily Uroctoninae
        • Family Euscorpiidae Laurie, 1896
          • Subfamily Euscorpiinae Laurie, 1896
          • Subfamily Megacorminae Kraepelin, 1905
            • Tribe Chactopsini Soleglad et Sissom, 2001
            • Tribe Megacormini Kraepelin, 1905
          • Subfamily Scorpiopinae Kraepelin, 1905
            • Tribe Scorpiopini Kraepelin, 1905
            • Tribe Troglocormini Soleglad et Sissom, 2001
        • Family Superstitioniidae Stahnke, 1940
          • Subfamily Superstitioniinae Stahnke, 1940
          • Subfamily Typlochactinae Mitchell, 1971
        • Family Vaejovidae Thorell, 1876
      • Superfamily Iuroidea Thorell, 1876
        • Family Iuridae Thorell, 1876
        • Family Caraboctonidae Kraepelin, 1905 (hairy scorpions)
          • Subfamily Caraboctoninae Kraepelin, 1905
          • Subfamily Hadrurinae Stahnke, 1974
      • Superfamily Scorpionoidea Latreille, 1802
        • Family Bothriuridae Simon, 1880
          • Subfamily Bothriurinae Simon, 1880
          • Subfamily Lisposominae Lawrence, 1928
        • Family Diplocentridae Karsch, 1880
        • Family Scorpionidae Latreille, 1802 (burrowing scorpions or pale-legged scorpions)
          • Subfamily Diplocentrinae Karsch, 1880
            • Tribe Diplocentrini Karsch, 1880
              • Tribe Nebini Kraepelin, 1905
          • Subfamily Scorpioninae Latreille, 1802
          • Subfamily Urodacinae Pocock, 1893
        • Family Hemiscorpiidae Pocock, 1893 (= Ischnuridae, =Liochelidae) (rock scorpions, creeping scorpions, or tree scorpions)
          • Subfamily Hemiscorpiinae Pocock, 1893
          • Subfamily Heteroscorpioninae Kraepelin, 1905
          • Subfamily Hormurinae Laurie, 1896

[edit] Cultural symbolism

The scorpion has had various meanings and representations in different cultures in history:

• In the Epic of Gilgamesh, Gilgamesh approaches mountains where scorpion-folk guard the entrance. Additionally, the Akkadians called the constellation Scorpius, Girtab, meaning "the Seizer", or "Stinger" and "Place Where One Bows Down".
• In ancient Egypt, the scorpion was associated with the god Set.
• The Falaknuma Palace of Hyderabad, India, is laid out in the shape of a scorpion with the two pincers spreading out to the north as wings to the building.
• In Greek mythology, the scorpion is conjured by the gods to hound and punish Orion. It is also said that when Perseus slew Medusa, the blood that leaked out of her severed neck turned into scorpions and snakes as it hit the ground.
• From a Biblical quotation, it is the term for a severe Roman scourge. Hard material was fixed onto multiple thongs to give them a flesh-tearing bite [1 Kings 12:11: ...My father scourged you with whips; I will scourge you with scorpions]. The choice of the name testifies how much the hellish pain caused by the small animal is to be feared.
• The Persian legendary monster manticore is often depicted with a scorpion tail.
• The scorpion is one of the symbols of the Astrological sign of Scorpio.
• The scorpion is the symbol of the Mexican state of Durango.

[edit] See also

• List of some notable scorpion species

Several species bear the name "scorpion" but do not belong to the order Scorpiones:

• List of false scorpions

[edit] References

Hadley NS & William SC (1968). Surface activity of some north american scoripions in relation to feeding. Ecology 49, 726-735. Stachel SJ, Stockwell SA, Van Vranken DL (1999). The fluorescence of scorpions and cataractogenesis. Chemistry and Biology 6, 531-539.

[edit] External links

• Information on scorpions and other arachnids
• Pepe the two-tailed scorpion
• Scorpion detection using UV LEDs (also movies of scorpions)
• Desert USA: Scorpions
• Photo gallery of several scorpions in captive breeding.
• Scorpion Venom Tested as Brain Cancer Treatment
• Scorpions as pets

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