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Scorpions are
predatory Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
arachnid Arachnida () is a class of joint-legged invertebrate animals ( arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and ...
s of the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
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 Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest co ...
. There are over 2,500 described
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
, with 22 extant (living) families recognized to date. Their
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
is being revised to account for 21st-century genomic studies. Scorpions primarily prey on
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
s and other
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s, but some species hunt
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s. 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
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
s harden, transporting them on her back. The exoskeleton contains
fluorescent Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, th ...
chemicals and glows under
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiati ...
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 In mathematics, a percentage (from la, per centum, "by a hundred") is a number or ratio expressed as a fraction of 100. It is often denoted using the percent sign, "%", although the abbreviations "pct.", "pct" and sometimes "pc" are also us ...
) 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 Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English ...
between 1175 and 1225 AD from
Old French Old French (, , ; Modern French: ) was the language spoken in most of the northern half of France from approximately the 8th to the 14th centuries. Rather than a unified language, Old French was a linkage of Romance dialects, mutually intelligi ...
', or from Italian ', both derived from the Latin ', equivalent to ', which is the
romanization Romanization or romanisation, in linguistics, is the conversion of text from a different writing system to the Roman (Latin) script, or a system for doing so. Methods of romanization include transliteration, for representing written text, an ...
of the Greek  – ', ultimately from
Proto-Indo-European root The roots of the reconstructed Proto-Indo-European language (PIE) are basic parts of words that carry a lexical meaning, so-called morphemes. PIE roots usually have verbal meaning like "to eat" or "to run". Roots never occurred alone in the la ...
''*(s)ker-'' meaning "to cut", cf. " shear".


Evolution


Fossil record

Scorpion
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s have been found in many
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
, including marine
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago (Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozoi ...
and estuarine
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, w ...
deposits, coal deposits from the Carboniferous Period and in
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In M ...
. Whether the early scorpions were marine or terrestrial has been debated, though they had
book lung A book lung is a type of respiration organ used for atmospheric gas exchange that is present in many arachnids, such as scorpions and spiders. Each of these organs is located inside an open ventral abdominal, air-filled cavity (atrium) and conn ...
s like modern terrestrial species. Over 100 fossil species of scorpion have been described. The oldest found as of 2021 is '' Dolichophonus loudonensis'', which lived during the Silurian, in present-day Scotland. ''
Gondwanascorpio ''Gondwanascorpio emzantsiensis'' is an extinct Gondwanan scorpion that lived 360 million years ago in the Devonian. Its fossil remains, clearly showing pincer and sting, were discovered in rocks of the Witteberg Group near Grahamstown in South A ...
'' from the Devonian is among the earliest-known terrestrial animals on the
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final st ...
supercontinent. Some Palaeozoic scorpions possessed
compound eye A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which dis ...
s similar to those of eurypterids. The
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
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.


Phylogeny

The Scorpiones are a clade within the pulmonate
Arachnida Arachnida () is a Class (biology), class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, came ...
(those with book lungs). Arachnida is placed within the Chelicerata, a subphylum of
Arthropoda Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
that contains sea spiders and horseshoe crabs, alongside terrestrial animals without book lungs such as ticks and
harvestmen The Opiliones (formerly Phalangida) are an Order (biology), order of arachnids Common name, colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs. , over 6,650 species of harvestmen have been discovered worldwide, alth ...
. The extinct Eurypterida, sometimes called sea scorpions, though they were not all marine, are not scorpions; their grasping pincers were
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as " jaws", chelicerae may be shaped as either articulated fangs, or simila ...
, not
homologous Homology may refer to: Sciences Biology *Homology (biology), any characteristic of biological organisms that is derived from a common ancestor *Sequence homology, biological homology between DNA, RNA, or protein sequences * Homologous chrom ...
with the pincers (second appendages) of scorpions. Scorpiones is sister to the Tetrapulmonata, a terrestrial group of pulmonates containing the
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species d ...
s and whip scorpions. This 2019
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
summarizes: Recent studies place
pseudoscorpion Pseudoscorpions, also known as false scorpions or book scorpions, are small, scorpion-like arachnids belonging to the order Pseudoscorpiones, also known as Pseudoscorpionida or Chelonethida. Pseudoscorpions are generally beneficial to humans sin ...
s as the sister group of scorpions in the clade Panscorpiones, which together with Tetrapulmonata makes up the clade Arachnopulmonata. The internal
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
of the scorpions has been debated, 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 Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
. The Iuroidea and Chactoidea are both seen not to be single clades, and are shown as "
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
" (with quotation marks) in this 2018 cladogram.


Taxonomy

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
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 ''Scorpio maurus'' is a species of North African and Middle Eastern scorpion, also known as the large-clawed scorpion or Israeli gold scorpion and lesser known as Zerachia scorpion. This is a small/medium-sized scorpion from the family Scorpion ...
'', '' Androctonus australis'', and ''
Euscorpius ''Euscorpius'' is a genus of scorpions, commonly called small wood-scorpions. It presently contains 65 species and is the type genus of the family Euscorpiidae – long included in the ChactidaeRein (2008a) – and the subfamily Euscorpiinae ...
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. In 1801,
Jean-Baptiste Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolog ...
divided up the "Insecta aptera", creating the
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
Arachnides for spiders, scorpions, and acari (mites and ticks), though it also contained the Thysanura (thrips), Myriapoda and parasites such as lice. 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". 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. There are over 100 described taxa of fossil scorpions. This classification is based on Soleglad and Fet (2003), which replaced Stockwell's older, unpublished classification. Further taxonomic changes are from papers by Soleglad et al. (2005). The extant taxa to the rank of
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
(numbers of species in parentheses) are: ; Order Scorpiones * Parvorder Pseudochactida Soleglad & Fet, 2003 ** Superfamily
Pseudochactoidea The Pseudochactidae are a scorpion family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of th ...
Gromov, 1998 *** Family
Pseudochactidae The Pseudochactidae are a scorpion family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of th ...
Gromov, 1998 (1 sp.) (Central Asian scorpions of semi- savanna habitats) * Parvorder Buthida Soleglad & Fet, 2003 ** Superfamily Buthoidea
C. L. Koch Carl Ludwig Koch (21 September 1778 – 23 August 1857) was a German entomologist and arachnologist. He was responsible for classifying a great number of spiders, including the Brazilian whiteknee tarantula and common house spider. He was born ...
, 1837
*** Family
Buthidae The Buthidae are the largest family of scorpions, containing about 100 genera and 1339 species as of 2022. A few very large genera ('' Ananteris'', '' Centruroides'', ''Compsobuthus'', or '' Tityus'') are known, but a high number of species-po ...
C. L. Koch Carl Ludwig Koch (21 September 1778 – 23 August 1857) was a German entomologist and arachnologist. He was responsible for classifying a great number of spiders, including the Brazilian whiteknee tarantula and common house spider. He was born ...
, 1837
(1209 spp.) (thick-tailed scorpions, including the most dangerous species) *** Family
Microcharmidae The Buthidae are the largest family of scorpions, containing about 100 genera and 1339 species as of 2022. A few very large genera (''Ananteris'', ''Centruroides'', ''Compsobuthus'', or '' Tityus'') are known, but a high number of species-poor or ...
Lourenço, 1996, 2019 (17 spp.) (African scorpions of humid forest leaf litter) * Parvorder Chaerilida Soleglad & Fet, 2003 ** Superfamily
Chaeriloidea ''Chaerilus'' is a genus of scorpions in the family Chaerilidae. They live in tropical parts of South Asia and Southeast Asia. A fossil genus '' Electrochaerilus'' is known from the Late Cretaceous (Cenomanian) aged Burmese amber. Description To ...
Pocock Pocock is a surname, and may refer to: *Andrew Pocock (born 1955), British High Commissioner to Nigeria * Barbara Pocock, one of the candidates of the 2022 Australian federal election * Bill Pocock (1884–1959), English footballer * Blair Pocock ...
, 1893
*** Family Chaerilidae
Pocock Pocock is a surname, and may refer to: *Andrew Pocock (born 1955), British High Commissioner to Nigeria * Barbara Pocock, one of the candidates of the 2022 Australian federal election * Bill Pocock (1884–1959), English footballer * Blair Pocock ...
, 1893
(51 spp.) (South and Southeast Asian scorpions of non-arid places) * Parvorder Iurida Soleglad & Fet, 2003 ** Superfamily Chactoidea
Pocock Pocock is a surname, and may refer to: *Andrew Pocock (born 1955), British High Commissioner to Nigeria * Barbara Pocock, one of the candidates of the 2022 Australian federal election * Bill Pocock (1884–1959), English footballer * Blair Pocock ...
, 1893
*** Family
Akravidae ''Akrav israchanani'' is an extinct species of scorpions from the Ayyalon Cave in Israel.Levy, 2007 "The first troglobite scorpion from Israel and a new chactoid family (Arachnida: Scorpiones)/ref> Description ''Akrav israchanani'' was an eyel ...
Levy, 2007 (1 sp.) (cave-dwelling scorpions of Israel) *** Family
Belisariidae Belisariidae is a family of scorpions belonging to the order Scorpiones. Genera There are two genera: * '' Belisarius'' Simon, 1879 * '' Sardoscorpius'' Tropea & Onnis, 2020 References Scorpions Scorpion families Taxa named by Wilson R. ...
Lourenço, 1998 (3 spp.) (cave-related scorpions of Southern Europe) *** Family
Chactidae The Chactidae are a family of scorpions established by Reginald Innes Pocock in 1893. They make up the Taxonomic rank, superfamily Chactoidea. Genera Chactidae contains the following genera: *''Anuroctonus'' Pocock, 1893 *''Auyantepuia'' (Gonz ...
Pocock Pocock is a surname, and may refer to: *Andrew Pocock (born 1955), British High Commissioner to Nigeria * Barbara Pocock, one of the candidates of the 2022 Australian federal election * Bill Pocock (1884–1959), English footballer * Blair Pocock ...
, 1893
(209 spp.) (New World scorpions, membership under revision) *** Family
Euscorpiidae The Euscorpiidae are a family of scorpions. Genera ''Euscorpiidae'' contains the following six genera: * '' Alpiscorpius'' Gantenbein ''et al.'', 1999 * ''Euscorpius ''Euscorpius'' is a genus of scorpions, commonly called small wood-scorpion ...
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 Typhlochactidae is a family of troglomorphic Troglomorphism is the morphological adaptation of an animal to living in the constant darkness of caves, characterised by features such as loss of pigment, reduced eyesight or blindness, and frequent ...
Mitchell, 1971 (11 spp.) (cave-related scorpions of Eastern Mexico) *** Family
Vaejovidae Vaejovidae is a family of scorpions, comprising 25 genera and 227 species, found in North America. The species of the genus are found in Mexico and the southern United States, and ''Paruroctonus boreus'' is found in Canada and is the northernmo ...
Thorell, 1876 (222 spp.) (New World scorpions) ** Superfamily Iuroidea Thorell, 1876 *** Family
Caraboctonidae The Caraboctonidae (hairy scorpions) are part of the superfamily Iuroidea. The family was established by Karl Kraepelin Karl Matthias Friedrich Magnus Kraepelin (; ; 14 December 1848 – 28 June 1915) was a German naturalist who specialised in ...
Kraepelin, 1905 (23 spp.) (hairy scorpions) *** Family
Hadruridae Hadruridae is a family of scorpions belonging to the order Scorpiones. Genera: * ''Hadrurus ''Hadrurus'' is a genus of scorpions which belongs to the family Caraboctonidae. They are found in sandy deserts and other xeric habitats in northwes ...
Stahnke, 1974 (9 spp.) (large North American scorpions) *** Family
Iuridae The Iuridae are a family of scorpions in the order Scorpiones. Six genera and at least 20 described species are placed in the Iuridae. Genera * '' Anuroctonus'' * ''Calchas'' * ''Hadrurus'' (giant hairy scorpions) * ''Iurus ''Iurus'' is a gen ...
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 ''Hemiscorpius'' is the sole genus of the scorpion family Hemiscorpiidae, with about 16 described species. Before Hemiscorpiidae, the term used for the family was Ischnuridae, which had to be changed due to a naming conflict with the damselfly fa ...
Pocock Pocock is a surname, and may refer to: *Andrew Pocock (born 1955), British High Commissioner to Nigeria * Barbara Pocock, one of the candidates of the 2022 Australian federal election * Bill Pocock (1884–1959), English footballer * Blair 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 ''Rugodentus'' is a monotypic genus of scorpions 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 charac ...
Bastawade et al., 2005 (1 sp.) (burrowing scorpions of India) *** Family Scorpionidae Latreille, 1802 (183 spp.) (burrowing or pale-legged scorpions) *** Family
Diplocentridae Diplocentridae is a family of scorpions. The roughly 120 species are mostly native to the New World, except for genus '' Nebo'', which is distributed in the Middle East.Santibáñez-López, C. E.; Francke, O. F.; and Prendini, L. (2014)''Kolotl'' ...
Karsch, 1880 (134 spp.) (closely related to and sometimes placed in Scorpionidae, but have spine on telson) *** Family
Heteroscorpionidae ''Heteroscorpion'' is a genus of scorpions belonging to the monotypic family Heteroscorpionidae. The species of this genus are found in Madagascar. Species: *'' Heteroscorpion goodmani'' *'' Heteroscorpion kaii'' *'' Heteroscorpion kraepelin ...
Kraepelin, 1905 (6 spp.) (scorpions of Madagascar)


Geographical distribution

Scorpions are found on all continents except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest co ...
. 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 Great Britain is an island in the North Atlantic Ocean off the northwest coast of continental Europe. With an area of , it is the largest of the British Isles, the largest European island and the ninth-largest island in the world. It is ...
but were accidentally introduced by humans, and have now established a population.
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 List of islands of New Zealand, smaller islands. It is the ...
, and some of the islands in
Oceania Oceania (, , ) is a region, geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern Hemisphere, Eastern and Western Hemisphere, Western hemispheres, Oceania is estimated to have a land area of ...
, have in the past had small populations of introduced scorpions, but they were exterminated. Five colonies of ''
Euscorpius flavicaudis ''Tetratrichobothrius flavicaudis'', or the European yellow-tailed scorpion, is a small black scorpion with yellow-brown legs and tail (metasoma). Adults measure about long. It has relatively large, strong claws (pedipalps) and a short, thin tai ...
'' have established themselves since the late 19th century in
Sheerness Sheerness () is a town and civil parish beside the mouth of the River Medway on the north-west corner of the Isle of Sheppey in north Kent, England. With a population of 11,938, it is the second largest town on the island after the nearby tow ...
in England at 51°N, while ''
Paruroctonus boreus ''Paruroctonus boreus'', commonly known as the ''northern scorpion'', is a species of scorpion in the family Vaejovidae. Ranging from Arizona to Canada. Distribution Can be as far north as 52° N. Several anecdotal reports show them found ne ...
'' lives as far north as
Red Deer, Alberta Red Deer is a city in Alberta, Canada, located midway on the Calgary–Edmonton Corridor. Red Deer serves central Alberta, and key industries include health care, retail trade, construction, oil and gas, hospitality, manufacturing and education. ...
, at 52°N. A few species are on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
; ''
Lychas braueri ''Afrolychas braueri'', commonly known as the Seychelles forest scorpion, is a species of scorpion in the family Buthidae. It is currently thought to survive only on Silhouette Island, Seychelles, although the species was historically found on ...
'' is classed as critically endangered (2014), ''
Isometrus deharvengi ''Isometrus'' is a genus of scorpion belonging and being eponymous to the family Buthidae. Some species are currently assigned to the genus ''Reddyanus''. Distribution The species of this genus are found in south and southeast Asia and in Oceani ...
'' as
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...
(2016) and '' Chiromachus ochropus'' as
vulnerable Vulnerable may refer to: General * Vulnerability * Vulnerability (computing) * Vulnerable adult * Vulnerable species Music Albums * ''Vulnerable'' (Marvin Gaye album), 1997 * ''Vulnerable'' (Tricky album), 2003 * ''Vulnerable'' (The Used album) ...
(2014). Scorpions are xerocoles, meaning they primarily live in deserts, but they can be found in virtually every terrestrial
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
including high-elevation mountains, caves, and
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...
s. They are largely absent from boreal ecosystems such as the
tundra In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless mo ...
, high-altitude
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruces ...
, and mountain tops. The highest altitude reached by a scorpion is in the Andes, for '' Orobothriurus crassimanus''. As regards microhabitats, scorpions may be ground-dwelling, tree-loving, rock-loving or sand-loving. Some species, such as ''
Vaejovis janssi ''Vaejovis janssi'' is a species of scorpion endemic to the Revillagigedo Islands in Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the nor ...
'', are versatile and are found in all habitats on Socorro Island,
Baja California Baja California (; 'Lower California'), officially the Free and Sovereign State of Baja California ( es, Estado Libre y Soberano de Baja California), is a state in Mexico. It is the northernmost and westernmost of the 32 federal entities of Mex ...
, while others such as '' Euscorpius carpathicus'',
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
to the
littoral zone The littoral zone or nearshore is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely inundated), to coastal ...
of rivers in Romania, occupy specialized niches.


Morphology

Scorpions range in size from the ''
Typhlochactas mitchelli ''Typhlochactas mitchelli'' is a species of scorpion of the family Typhlochactidae. It is endemic to the state of Oaxaca, Mexico. This species is of the eyeless cave-dwelling genus '' Typhlochactas''. References External links "Phylogenetic ...
'' of Typhlochactidae, to the ''
Heterometrus swammerdami ''Gigantometrus swammerdami'', commonly called the giant forest scorpion, is a scorpion belonging to the family Scorpionidae. It is native to India. It is the world's largest scorpion species with 23 cm (9 in) in length, and weigh as mu ...
'' of Scorpionidae. The body of a scorpion is divided into two parts or tagmata: the
cephalothorax The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''ceph ...
or prosoma, and the abdomen or
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma ( cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects ...
. 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. External differences between the sexes are not obvious in most species. In some, the metasoma is more elongated in males than females.


Cephalothorax

The cephalothorax comprises the
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the und ...
, eyes, chelicerae (mouth parts), pedipalps (which have
chelae A chela ()also called a claw, nipper, or pinceris a pincer-like organ at the end of certain limbs of some arthropods. The name comes from Ancient Greek , through New Latin '. The plural form is chelae. Legs bearing a chela are called chelipeds. ...
, commonly called claws or pincers) and four pairs of
walking leg The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip (anat ...
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. The chelicerae are at the front and underneath the carapace. They are pincer-like and have three segments and sharp "teeth". The brain of a scorpion is in the back of the cephalothorax, just above the
esophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
. 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 Primitive may refer to: Mathematics * Primitive element (field theory) * Primitive element (finite field) * Primitive cell (crystallography) * Primitive notion, axiomatic systems * Primitive polynomial (disambiguation), one of two concepts * Pr ...
trait. The pedipalp is a segmented, clawed
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including ant ...
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. 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. Depending on the species, the legs may have spines and spurs.


Mesosoma

The mesosoma or preabdomen is the broad part of the opisthosoma. 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. 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, which function as sensory organs. 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. 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. The 7th and last somite does not bear appendages or any other significant external structures. 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
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces ...
s 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 An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The femal ...
formation and embryonic development. They connect to two
oviduct The oviduct in mammals, is the passageway from an ovary. In human females this is more usually known as the Fallopian tube or uterine tube. The eggs travel along the oviduct. These eggs will either be fertilized by spermatozoa to become a zygote, ...
s which connect to a single atrium leading to the genital orifice. Males have two gonads made of two cylindrical tubes with a ladder-like configuration; they contain cysts which produce
spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell (biology), cell, or moving form of the ploidy, haploid cell (biology), cell that is the male gamete. A spermatozoon Fertilization, joins an ovum to form a zygote. (A zygote ...
. Both tubes end in a
spermiduct The ejaculatory ducts (''ductus ejaculatorii'') are paired structures in male anatomy. Each ejaculatory duct is formed by the union of the vas deferens with the duct of the seminal vesicle. They pass through the prostate, and open into the ureth ...
, 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 Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
-based structures which come together to form the spermatophore.


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. The tails of some species contain light receptors. The telson includes the vesicle, which contains a symmetrical pair of
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a ...
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. 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. The stinger contains metalloproteins with zinc, hardening the tip. The optimal stinging angle is around 30 degrees relative to the tip.


Biology

Most scorpion species are
nocturnal Nocturnality is an ethology, animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatures generally have ...
or
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of dayli ...
, finding shelter during the day in burrows, cracks in rocks and tree bark. 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 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. Scorpions prefer areas where the temperature remains in the range of , but may survive temperatures from well below freezing to desert heat. Scorpions can withstand intense heat: ''
Leiurus quinquestriatus The deathstalker (''Leiurus quinquestriatus'') is a species of scorpion, a member of the family Buthidae. It is also known as the Palestine yellow scorpion, Omdurman scorpion, Naqab desert scorpion and by many other colloquial names, which gen ...
'', ''Scorpio maurus'' and '' Hadrurus arizonensis'' can live in temperatures of 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 . 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 Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
. 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. Desert scorpions have several adaptations for water conservation. They excrete insoluble compounds such as
xanthine Xanthine ( or ; archaically xanthic acid; systematic name 3,7-dihydropurine-2,6-dione) is a purine base found in most human body tissues and fluids, as well as in other organisms. Several stimulants are derived from xanthine, including caffeine, ...
,
guanine Guanine () (symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine ( uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is c ...
, and
uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the metabolic breakdown ...
, 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 Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids incl ...
s and waxes from epidermal glands, and protects against ultraviolet radiation. Even when dehydrated, a scorpion can tolerate high osmotic pressure in its blood. 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. 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 in a defensive strike.


Mortality and defense

Scorpions may be attacked by other arthropods like ants, spiders, solifugids and centipedes. Major
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
s include frogs, lizards, snakes, birds, and mammals. Meerkats are somewhat specialized in preying on scorpions, biting off their stingers and being immune to their venom. Other predators adapted for hunting scorpions include the
grasshopper mouse Grasshopper mice are rodents of the genus ''Onychomys'', occurring in North America. They feed on insects and other arthropods. Biology The three species in this genus of New World mice are only distantly related to the common house mouse, '' ...
and
desert long-eared bat The desert long-eared bat (''Otonycteris hemprichii'') is a species of vesper bat found in North Africa and the Middle East. Taxonomy The taxonomic classification of ''Otonycteris'' is unclear, however their chromosomes suggest that this genus i ...
, which are also immune to their venom. In one study, 70% of the latter's droppings contained scorpion fragments. Scorpions host parasites including
mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
s,
scuttle flies The fly ''Megaselia scalaris'' is a member of the order Diptera and the family Phoridae, and it is widely distributed in warm regions of the world. The family members are commonly known as the "humpbacked fly", the "coffin fly", and the "scuttle ...
, nematodes and some bacteria. The
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as Tumor immunology, cancer cells and objects such ...
of scorpions gives them resistance to infection by many types of bacteria. When threatened, a scorpion raises its claws and tail in a defensive posture. Some species
stridulate Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
to warn off predators by rubbing certain hairs, the stinger or the claws. Certain species have a preference for using either the claws or stinger as defense, depending on the size of the appendages. A few scorpions, such as '' Parabuthus'', '' Centruroides margaritatus'', and ''Hadrurus arizonensis'', squirt venom in a narrow jet as far as to warn off potential predators, possibly injuring them in the eyes. 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.


Diet and feeding

Scorpions generally prey on insects, particularly grasshoppers,
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by st ...
s,
termite Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...
s,
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s and wasps. Other prey include spiders,
solifugid Solifugae is an order of animals in the class Arachnida known variously as camel spiders, wind scorpions, sun spiders, or solifuges. The order includes more than 1,000 described species in about 147 genera. Despite the common names, they are nei ...
s, woodlice and even small
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
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 ''Isometroides vescus'', also known as the spider-hunting scorpion or spiral burrow scorpion, is a species of scorpion in the Buthidae family. It is native to Australia, and was first described by German arachnologist Ferdinand Karsch in 1880. ...
'' 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 A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, a ...
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. 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 An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
fragments) is trapped by setae in the pre-oral cavity and ejected. The sucked-in food is pumped into the midgut by the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its ...
, 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 Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cel ...
, and a relatively inactive lifestyle. This enables some to survive six to twelve months of starvation.


Mating

Most scorpions reproduce sexually, with male and female individuals; species in some genera, such as ''
Hottentotta ''Hottentotta'' is a genus of scorpions of the family Buthidae. It is distributed widely across Africa, except for most of the Sahara desert. Species in the genus also occur in the Middle East, the Arabian Peninsula, southeastern Turkey, Iraq, I ...
'' and '' Tityus'', and the species ''
Centruroides gracilis ''Centruroides gracilis'' is a species of scorpion in the family Buthidae, the bark scorpions. Its common names include Florida bark scorpion, brown bark scorpion, and slender brown scorpion.Rein, J. O''Centruroides gracilis''.The Scorpion Files. ...
'', '' Liocheles australasiae'', and '' Ananteris coineaui'' have been reported, not necessarily reliably, to reproduce through
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
, in which unfertilized eggs develop into living
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm ...
s. 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. 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. 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. Sexual cannibalism after mating has only been reported anecdotally in scorpions.


Birth and development

Gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during preg ...
in scorpions can last for over a year in some species. They have two types of
embryonic development An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
; 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 though. Unlike the majority of arachnids, which are
oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), a ...
, hatching from eggs, scorpions seem to be universally viviparous, with live births. They are unusual among terrestrial arthropods in the amount of care a female gives to her offspring. The size of a brood varies by species, from 3 to over 100. The body size of scorpions is not correlated either with brood size or with life cycle length. 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. 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. 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.


Fluorescence

Scorpions glow a vibrant blue-green when exposed to certain wavelengths ranges of
ultraviolet Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiati ...
light such as that produced by a
black light A blacklight, also called a UV-A light, Wood's lamp, or ultraviolet light, is a lamp that emits long-wave ( UV-A) ultraviolet light and very little visible light. One type of lamp has a violet filter material, either on the bulb or in a se ...
, due to
fluorescent Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, th ...
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. This fluorescence may have an active role in the scorpion's ability to detect light.


Relationship with humans


Stings

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. People with allergies are especially at risk; otherwise, first aid is symptomatic, with
analgesia Pain management is an aspect of medicine and health care involving relief of pain (pain relief, analgesia, pain control) in various dimensions, from acute and simple to chronic and challenging. Most physicians and other health professionals ...
. Cases of very high blood pressure are treated with medications that relieve anxiety and relax the blood vessels. 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
benzodiazepine Benzodiazepines (BZD, BDZ, BZs), sometimes called "benzos", are a class of depressant drugs whose core chemical structure is the fusion of a benzene ring and a diazepine ring. They are prescribed to treat conditions such as anxiety disorders, ...
s when there is neuromuscular involvement. Although rare, severe
hypersensitivity reaction Hypersensitivity (also called hypersensitivity reaction or intolerance) refers to undesirable reactions produced by the normal immune system, including allergies and autoimmunity. They are usually referred to as an over-reaction of the immune ...
s including
anaphylaxis Anaphylaxis is a serious, potentially fatal allergic reaction and medical emergency that is rapid in onset and requires immediate medical attention regardless of use of emergency medication on site. It typically causes more than one of the follo ...
to scorpion antivenin are possible. 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. Mexico is one of the most affected countries, with the highest
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity' ...
of scorpions in the world, some 200,000 envenomations per year and at least 300 deaths. 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.


Potential medicinal use

Scorpion venom is a mixture of neurotoxins; most of these are
peptide Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. ...
s, chains of
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
s. Many of them interfere with membrane channels that transport sodium,
potassium Potassium is the chemical element with the symbol K (from Neo-Latin '' kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosp ...
,
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...
, or
chloride The chloride ion is the anion (negatively charged ion) Cl−. It is formed when the element chlorine (a halogen) gains an electron or when a compound such as hydrogen chloride is dissolved in water or other polar solvents. Chloride s ...
ions. These channels are essential for
nerve conduction An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ca ...
,
muscle contraction Muscle contraction is the activation of tension-generating sites within muscle cells. In physiology, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in muscle length, such as ...
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 An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
, 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.


Consumption

Scorpions are eaten by people in West Africa, Myanmar and East Asia. Fried scorpion is traditionally eaten in Shandong, China. 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. In Thailand, scorpions are not eaten as often as other arthropods, such as grasshoppers, but they are sometimes fried as street food. They are used in Vietnam to make snake wine (scorpion wine).


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 Peat (), also known as turf (), is an accumulation of partially decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, moors, or muskegs. The peatland ecosystem covers and is the most efficien ...
for forest species, or
lateritic Laterite is both a soil and a rock type rich in iron and aluminium and is commonly considered to have formed in hot and wet tropical areas. Nearly all laterites are of rusty-red coloration, because of high iron oxide content. They develop b ...
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 Cannibalism is the act of consuming another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded in more than 1,500 species. Human cannibalism is well documented, b ...
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. The pet trade has threatened wild populations of some scorpion species, particularly ''Androctonus australis'' and ''
Pandinus imperator The emperor scorpion, ''Pandinus imperator'', is a species of scorpion native to rainforests and savannas in West Africa. It is one of the largest scorpions in the world and lives for 6–8 years. Its body is black, but like other scorpions it ...
''.


Culture

File:Egyptian - Figure of Isis-Serget as Scorpion - Walters 54546 - Side A (cropped).jpg, Late period bronze figure of Isis-
Serket Serket ( egy, , italics=no, translit=srqt) is the goddess of healing venomous stings and bites in Egyptian mythology, originally the deification of the scorpion. Her family life is unknown, but she is sometimes credited as the daughter of Neith ...
Scorpion and snake fighting Anglo-Saxon c 1050.jpg, "Scorpion and snake fighting", ''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 scorpion motif (two types shown) was often woven into Turkish '' kilim'' flatweave carpets, for protection from their sting.
The scorpion is a culturally significant animal, appearing as a
motif Motif may refer to: General concepts * Motif (chess composition), an element of a move in the consideration of its purpose * Motif (folkloristics), a recurring element that creates recognizable patterns in folklore and folk-art traditions * Moti ...
in art, especially in
Islamic art Islamic art is a part of Islamic culture and encompasses the visual arts produced since the 7th century CE by people who lived within territories inhabited or ruled by Muslim populations. Referring to characteristic traditions across a wide r ...
in the Middle East. A scorpion motif is often woven into Turkish kilim flatweave carpets, for protection from their sting. The scorpion is perceived both as an embodiment of evil and as a protective force such as a dervish's powers to combat evil. In Muslim folklore, the scorpion portrays
human sexuality Human sexuality is the way people experience and express themselves sexually. This involves biological, psychological, physical, erotic, emotional, social, or spiritual feelings and behaviors. Because it is a broad term, which has varied w ...
. Scorpions are used in
folk medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
in South Asia, especially in antidotes for scorpion stings. 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 Babylonian astronomy was the study or recording of celestial objects during the early history of Mesopotamia. Babylonian astronomy seemed to have focused on a select group of stars and constellations known as Ziqpu stars. These constellations ...
during the Chaldean period. This was then taken up by western
astrology Astrology is a range of divinatory practices, recognized as pseudoscientific since the 18th century, that claim to discern information about human affairs and terrestrial events by studying the apparent positions of celestial objects. Di ...
; in astronomy the corresponding constellation is named Scorpius. In ancient Egypt, the goddess
Serket Serket ( egy, , italics=no, translit=srqt) is the goddess of healing venomous stings and bites in Egyptian mythology, originally the deification of the scorpion. Her family life is unknown, but she is sometimes credited as the daughter of Neith ...
, who protected the
Pharaoh Pharaoh (, ; Egyptian: '' pr ꜥꜣ''; cop, , Pǝrro; Biblical Hebrew: ''Parʿō'') is the vernacular term often used by modern authors for the kings of ancient Egypt who ruled as monarchs from the First Dynasty (c. 3150 BC) until th ...
, was often depicted as a scorpion. In
ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
, a warrior's shield sometimes carried a scorpion device, as seen in
red-figure pottery Red-figure vase painting is one of the most important styles of figural Greek vase painting. It developed in Athens around 520 BCE and remained in use until the late 3rd century BCE. It replaced the previously dominant style of black-figure va ...
from the 5th century BC. In
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities of ...
,
Artemis In ancient Greek mythology and Ancient Greek religion, religion, Artemis (; grc-gre, Ἄρτεμις) is the goddess of the hunt, the wilderness, wild animals, nature, vegetation, childbirth, Kourotrophos, care of children, and chastity. ...
or
Gaia In Greek mythology, Gaia (; from Ancient Greek , a poetical form of , 'land' or 'earth'),, , . also spelled Gaea , is the personification of the Earth and one of the Greek primordial deities. Gaia is the ancestral mother—sometimes parthe ...
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. Scorpions are mentioned in the
Bible The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts o ...
and the
Talmud The Talmud (; he, , Talmūḏ) is the central text of Rabbinic Judaism and the primary source of Jewish religious law ('' halakha'') and Jewish theology. Until the advent of modernity, in nearly all Jewish communities, the Talmud was the cen ...
as symbols of danger and maliciousness. The
fable Fable is a literary genre: a succinct fictional story, in prose or verse, that features animals, legendary creatures, plants, inanimate objects, or forces of nature that are anthropomorphized, and that illustrates or leads to a particular mor ...
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. 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. Scorpions have equally appeared in western artforms including film and poetry: the
surrealist Surrealism is a cultural movement that developed in Europe in the aftermath of World War I in which artists depicted unnerving, illogical scenes and developed techniques to allow the unconscious mind to express itself. Its aim was, according to ...
filmmaker Luis Buñuel made symbolic use of scorpions in his 1930 classic ''
L'Age d'or ''L'Age d'Or'' (french: L'Âge d'Or, ), commonly translated as ''The Golden Age'' or ''Age of Gold'', is a 1930 French surrealist satirical comedy film directed by Luis Buñuel about the insanities of modern life, the hypocrisy of the sexual ...
'' (''The Golden Age''), while Stevie Smith's last collection of poems was entitled ''Scorpion and other Poems''. A variety of martial arts films and video games have been entitled ''Scorpion King''. Since classical times, the scorpion with its powerful stinger has been used to provide a name for weapons. In the
Roman army The Roman army (Latin: ) was the armed forces deployed by the Romans throughout the duration of Ancient Rome, from the Roman Kingdom (c. 500 BC) to the Roman Republic (500–31 BC) and the Roman Empire (31 BC–395 AD), and its medieval contin ...
, the scorpio was a torsion siege engine used to shoot a projectile.
Vitruvius Vitruvius (; c. 80–70 BC – after c. 15 BC) was a Roman architect and engineer during the 1st century BC, known for his multi-volume work entitled '' De architectura''. He originated the idea that all buildings should have three attribut ...
, ''De Architectura'', X:10:1–6.
The British Army's
FV101 Scorpion The FV101 Scorpion is a British armoured reconnaissance vehicle, and also a light tank. It was the lead vehicle and the fire support type in the Combat Vehicle Reconnaissance (Tracked), CVR(T), family of seven armoured vehicles. Manufactured b ...
was an armored reconnaissance vehicle or light
tank A tank is an armoured fighting vehicle intended as a primary offensive weapon in front-line ground combat. Tank designs are a balance of heavy firepower, strong armour, and good battlefield mobility provided by tracks and a powerful e ...
in service from 1972 to 1994. A version of the Matilda II tank, fitted with a flail to clear mines, was named the Matilda Scorpion. Several ships of the Royal Navy and of the US Navy have been named ''Scorpion'' including an 18-gun sloop in 1803, a turret ship in 1863, a patrol yacht in 1898, a destroyer in 1910, and a nuclear submarine in 1960. The scorpion has served as the name or symbol of products and brands including Italy's Abarth racing cars and a Montesa scrambler
motorcycle A motorcycle (motorbike, bike, or trike (if three-wheeled)) is a two or three-wheeled motor vehicle steered by a handlebar. Motorcycle design varies greatly to suit a range of different purposes: long-distance travel, commuting, cruisin ...
. 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.


Notes


References


Sources

* * *


External links


American Museum of Natural History - Scorpion Systematics Research Group

CDC – Insects and Scorpions – NIOSH Workplace Safety and Health Topic
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