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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 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.

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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]

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]

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]

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]

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 51N,[29][30][31] while Paruroctonus boreus lives as far north as Red Deer, Alberta, at 52N.[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]

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]

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]

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] 17dc91bb1f