Scorpions
| Scorpions | ||||||||||||
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Androctonus crassicauda |
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| Scorpiones | ||||||||||||
| CL Koch , 1837 |
The scorpions (Scorpiones) are an order of the arachnids (Arachnida). Depending on the classification, between 1750 and 2500 species are known worldwide, of which only about 25 are considered to be potentially fatal to humans. Scorpions reach body sizes between 9 millimeters in the species Typhlochactas mitchelli and 21 centimeters in the imperial scorpions ( Pandinus imperator ) and Hadogenes troglodytes. They live mainly in sandy or stony soils or near the bottom of the tropics and subtropics , deserts and semi-deserts . Few species are climbing tree-dwellers, hikers or cave dwellers and stay as cultural followers in the vicinity of human dwellings.
Den of the scorpions
External anatomy
The body of the scorpion is indistinctly divided into a front body ( prosoma ) and a clearly two-part abdomen (opisthosoma). The opisthosoma consists of a broad part, the mesosoma, and a tail-like elongated metasoma.
The front body consists of six segments and carries the extremities. They include the relatively small, three-part jaw claws ( Chelicere ), which are followed by the imposing Pedipalps . These are formed into large tentacles that end in scissors. Scorpions are able to dig burrows and burrows with their powerful claws. The scissors are also used to catch and hold on to prey, mostly other arthropods or smaller vertebrates . The chelicerae serve to chop up food and work against the bases of the pedipalps and the following two pairs of legs, which together form the lower boundary of the oral cavity (gnatho base).
The scissor-studded pedipalps are followed by four pairs of legs. The second abdomen segment of the scorpion carries the genital plates and in the rear area conspicuous comb-like structures, which are called comb organ (pecten). In the abdomen, like the spider , sits the fan lung . The anus lies in the fifth abdomen segment.
The abdomen, the opisthosoma, consists of chitin rings that are flexibly connected to one another. As a result, they are dimensionally stable and at the same time have extreme mobility. When moving, this part is carried upright over the body of the scorpion. The opisthosoma has 13 segments, of which the last 5 segments are narrowed to narrow rings as in the sea scorpions (Eurypterids) and form the metasoma. This metasoma carries the telson (terminal spine and poisonous bladder). Larger prey animals are killed with a sting through the stinger .
The scorpions perceive their prey mainly through their cleft sensory organs . A scorpion recognizes a digging cockroach by its vibrations from 50 centimeters away. The eyes of the scorpions are only suitable for rough orientation (position of the sun, moonlight, etc.). This nervous performance is provided by a chain of ganglia , the abdominal ganglion chain , which in weaving spiders has already fused to form the abdominal ganglion. The front body is also equipped with a large medial pair of eyes and up to five smaller pairs of point eyes .
The cuticle of the scorpion fluoresces when exposed to ultraviolet radiation . This stimulates stored beta-carbolines and 7-hydroxy-4-methylcoumarin. With the help of appropriate lamps, the animals can therefore be easily spotted in the dark. This effect is retained even after the animals die.
Internal anatomy
As with all arthropods, the muscles of the scorpions attach to the inner surfaces and clasps of the chitin skeleton . They are innervated by nerves radiating from a central abdominal marrow with seven nerve nodes (ganglia). In addition to the abdominal marrow, there is also a brain made up of two large ganglia, which are located in the head area and span the throat area .
The digestive system begins with a mouth area, which is equipped with a muscular throat. This works like a pump that sucks the predigested food into the mouth, from where it is then directed into the foregut and midgut. Digestion takes place in the midgut, into which several glands open which produce the necessary enzymes such as amylases , proteases and lipases . The storage organ is a large hepatopancreas (equivalent to a combination of liver and pancreas ), which can make up up to 20 percent of the animals' body weight. The nutrients are stored as glycogen . The metabolism rate of the animals is very low, and the food can be used very effectively. In addition, scorpions can ingest up to a third of their own body weight with food. These adaptations allow Scorpios to starve for up to 12 months.
The excretion takes place, as with other arthropods also about Malpighian vessels , which end in the intestine in the transition area between the midgut and hindgut and nitrogen compounds give into it. This excretion takes place with very little water loss, the substances are released as uric acid with the feces .
The vascular system is open with the exception of the dorsal cardiac tube, blood ( hemolymph ) floats freely in the body and corresponding in the blood sinus in the tissue of animals. The breathing takes place via book lungs that are present on the underside of scorpions as tucking of the cuticle. In these the oxygen is released into the hemolymph.
In both sexes, the gonads are paired as a network of tubes, which cannot be distinguished at first glance. The males produce sperm in their testicular tubes , which are packed into spermatophores in special organs (paraxial organs) . The females produce eggs that are laid out with or without a yolk supply depending on the species . With the apoicogenic eggs there is a yolk that is used by the embryos as food, the young scorpions from katoicogenic eggs instead use their chelicerae to grab special food diverticula in the female uterus and are nourished in this way.
distribution and habitat
Scorpions can be found worldwide on all continents with the exception of Antarctica . In America their distribution area extends from southern Canada to southern South America. In Europe you can find them with a northern distribution to the south of Austria and Switzerland. Scorpions were introduced as neozoa in Great Britain and New Zealand . However, the focus is on tropical and subtropical regions. The greatest biodiversity is found in the Mexican desert regions. The animals are found in most habitats such as deserts and semi-deserts, in the grassy savannah, in tropical forests, on coasts in the intertidal zone, and occasionally in caves. Many burrow underground, while some species live in trees. Alacran tartarus is a cave dweller and can be found up to 800 meters below the surface.
Most species live on the ground and, according to McDaniels 1968, are divided into four basic types:
- Psammophilic scorpions are adapted to sandy habitats. They are very fast runners on this surface and are well protected against dehydration.
- Lithophilic scorpions prefer to live in rock habitats and are usually built flat so that they can move easily between stones.
- Burrowing scorpions live mostly underground in caves they have dug themselves. They only leave them for hunting and reproduction.
- Wandering scorpions change their habitat and are therefore not very well adapted to certain conditions.
Scorpions are traditionally considered to thrive in dry habitats, but many species are dependent on high humidity.
Way of life
nutrition
Scorpions feed on various invertebrates such as insects (Insecta) or arachnids (Arachnida), and more rarely on snails or small vertebrates such as rodents , snakes and lizards . The choice of food is more or less specific to the species, only the Isometroides vescus species is considered a food specialist and feeds exclusively on a few burrowing spider species.
Scorpions are mostly nocturnal. Most species lie in wait for their prey in the vicinity of their burrows or other hiding places. Some species are also able to catch flying insects. There are also active hunters among the scorpions, these are mostly slender animals with very strong scorpion poisons . When catching prey, the two pedipalps are used as grippers; if the victim cannot be rendered incapable of defending himself, the sting is used, which is passed over the head in less than a second and injects poison into the prey. It is now crushed with the jaw claws, while enzymes pre-digest the food at the same time. The liquid food pulp is then sucked through the throat into the intestine. The eating process can take several hours.
Many scorpions are able to get along without food for a long time, some species even for one to two years, because their resting metabolism hardly requires any energy.
Predators and Defense
Scorpions are popular prey for various birds (especially owls ), lizards, snakes, large frogs, and mammals. In addition, many scorpions hunt other scorpions themselves, both native and other species, and even among larger arthropods they have enemies such as centipedes , roller spiders, and spiders .
In order to defend themselves against predators, scorpions have several defense mechanisms. The most striking defense is the equipment with a venomous sting and a highly potent venom , which is produced in the venom gland in the last segment of the body and consists of two components in most scorpions: one for killing arthropods as food and one against vertebrates for defense. Depending on the species, the venom has either a neuromuscular effect (including Centruroides spp. And Parabuthus spp. ) Or cardiovascular (including Buthus spp. , Mesobuthus spp. And Androctonus spp. ). The neurotoxic components are particularly relevant for the poisoning process itself .
The venom of most scorpions is usually not very dangerous to large vertebrates. As mentioned in the introduction, however, a few species have poisons that can lead to potentially fatal cardiac and central nervous symptoms in humans . This affects around 10% of all stab accidents, with older people, (small) children and the sick being particularly at risk. Regions in which there are frequent (fatal) accidents include Mexico and Iran. Particularly strong poisons are found in representatives of the Buthidae ( LD 50 value between 0.25 ppm (parts in a million) and 4.25 ppm in the mouse). Every year around 1000 to 5000 people die from scorpion stings worldwide, depending on the source.
Some species have specialized in scorpions to the point that they are immune to the poisons or have behaviors that enable them to avoid the sting. Their nocturnal way of life and their behavior of almost always hiding also avoids contact with potential enemies.
Social behavior
Most scorpions are solitary animals who only meet other scorpions in the postpartum period, when they are hunters or hunted, and during the mating season.
However, some species show pronounced social behavior. There are species that winter together in the same shelter and form aggregations for this purpose. With some species, such as the imperial scorpion ( Pandinus imperator ), the bond between the young scorpions of a litter remains and they form family groups that even hunt together.
Reproduction and development
One of the most important prerequisites for the settlement of the extremely dry habitat is the guarantee of reproduction and thus the protection of the eggs and sperm from dehydration .
Mating and mating dance
The males of the scorpions place the sperm in a specially designed container, the spermatophore . This offers the sperm protection against external influences. However, since most species live in very dry areas, this protection alone is not sufficient if the spermatophore is not ingested by the female within a very short time. The “wedding dance” of the scorpions serves this function. One such was shown in the Walt Disney film The Desert Lives .
During the mating season, the females exude sexual attractants ( pheromones ) that lead the males to them. Once the males have found one, they try to put it in a mating mood by twitching (juddering). Once the male has "persuaded" his partner, they grab the scissors and the mating dance, which sometimes lasts for hours, begins. At the beginning of the mating dance, the males of many scorpion species do not only hold on to the female with claws. They stick their poison sting into the thin skin on the female's scissor arm. Stimulation of the female is suspected; however, it is not clear whether the male also injects poison into the female.
In the mating dance, the male sometimes leads the female over many meters and tries to find a suitable place for his spermatophore with the comb organs (pectines) on the belly side. Once it has felt it, it pauses briefly and deposits the spermatophore. Then the female is pulled over it so that the sperm can penetrate directly into her genital pore. This ends the dance and the partners quickly separate - sometimes it ends with the consumption of the husband ( cannibalism ). Both sexes can mate several times in their lifetime, whereby females of the Buthidae have also been observed during mating while they were still carrying the brood of the last mating on their backs.
development
After a few (up to twelve) months, the female gives birth to living young ( viviparia ), so the eggs are already "hatched" in the uterus . The number of young scorpions can range from 2 to over 100, depending on the species. The young scorpions are white at birth, and each is individually enclosed by an embryo skin, the chorion . After the young scorpions have freed themselves from this, they climb onto the mother's back, who carries them around until the first molt. The first molt occurs after 1 to 51 days, depending on the species and external conditions. During this time the females are particularly aggressive. During this time, the young scorpions are fed by the body's own reserves, and they get fluids through the mother's back skin.
After the first molt, the young leave their mother and are on their own. The further development runs over several, usually five to eight, further moults. Then the animals are sexually mature. After reaching this sexual maturity, no further moulting takes place.
Danger
There are currently three types of scorpions on the Red List of the International Union for Conservation of Nature and Natural Resources (IUCN): Isometrus deharvengi , a höhlenbewohnender Scorpio from Vietnam, and Chiromachus ochropus and Lychas braueri , two endemic species of the Seychelles. Three species, the imperial scorpion ( Pandinus imperator ) as well as Pandinus dictator and Pandinus gambiensis are listed in Appendix II of the Washington Convention on Endangered Species and are subject to export controls.
Tribal history
As rural dwellers with a relatively thin layer of chitin , scorpions very rarely leave fossil remains, so little is known about the evolution of animals. Most of the knowledge comes from phylogenetic research . Due to the position of the scorpions at the base of the arachnids, it can be assumed that the scorpions are descended from marine forms, which at the same time may have been the parent species of the horseshoe crabs (Xiphosura) and the extinct sea scorpions (Eurypterida). All marine species still used gills for breathing, which in horseshoe crabs were placed on the inner back of the extremities. The scorpion's fan lungs emerged from the gills of their ancestors.
The first fossils of clearly terrestrial scorpions were found from the late Silurian about 430 to 390 million years ago. These early species were probably amphibious forms that were endowed with gills and adapted to life on the seashore and in the tidal area .
A splitting of the forms also began at this time and was already completed in the Devonian or at the latest in the Carboniferous about 325 million years ago. From this period, fossils of almost all scorpion taxa living today are known, the largest of which were more than 85 centimeters long.
The scorpions represent the most original group within the arachnids and are accordingly regarded as the sister group of all other arachnids.
Systematics
The systematic classification of the scorpions is based on morphological properties such as the number and distribution of the trichobothria , the shape of the sternum , the mouth structures, the teeth of the chelicerae, the shape of the legs, the telson and many other features. In addition, characteristics of embryology and internal anatomy play a role.
Currently, the ( recent ) scorpions living today are mostly divided into 13 families (according to Fet et al. 2000):
- Bothriuridae Simon , 1880
- Buthidae C.L. Koch , 1837
- Caraboctonidae Kraepelin , 1905
- Chactidae Pocock , 1893
- Chaerilidae Pocock , 1893
- Euscorpiidae Laurie , 1896
- Hemiscorpiidae Pocock , 1893
- Iuridae Thorell , 1876
- Microcharmidae Lourenço , 1996
- Pseudochactidae Gromov , 1998
- Scorpionidae Latreille , 1802
- Superstitioniidae Stahnke , 1940
- Vaejovidae Thorell , 1876
The Buthidae are the largest family of scorpions (Scorpiones) with more than 1,100 species in 90 genera . This family also includes most of the dangerously poisonous species. Another family, the Akravidae known only from Akrav israchanani , is believed to have died out in the second half of the 20th century.
Human and scorpions
Folk medicine
As an image, scorpions should avert the evil eye , which is why you can find their image on lamps and other objects. As a talisman , a scorpion is said to have protected the city of Emesa in the Middle East from real scorpions and snakes .
As a poisonous animal, the scorpion has found various uses in folk medicine. According to Megenberg, scorpion ashes, drunk in wine, and scorpion oil, with which one had to rub the stinging points, could help against the poison of the animals. The “scorpion oil” was obtained by dipping live scorpions in olive oil and then heating it. This oil should be able to heal a wide variety of ailments, including wounds , colic , gout and earache. In Tyrol , the oil was even used against urinary tract problems and the plague, as well as against bee and wasp stings . The bile of the scorpions was used against eye diseases and for skin beautification.
In superstition and folk medicine , the scorpion plays a major role in some peoples, mainly because of its poison. Konrad von Megenberg described the animals in his book of nature as
- “A species of snake which has a very delicate face, to be compared with the countenance of a chaste virgin. Those who are poisoned by the scorpion have three days before they have to die. "
He continues his description by telling of two-spined scorpions, which Aristotle reported. Among the pigs , the poison is only supposed to safely kill black pigs; the poison effect is accelerated when they go into the water. In humans, the scorpion only attacks hairy parts of the body and never prick the cupped hand. Konrad von Megenberg has another curiosity to report:
- “If you drown a scorpion in oil and pour vinegar on it in sunlight, it comes to life immediately. The oil clogs the small openings on his body, which in humans are called sweat holes and in Latin pori. The vinegar, on the other hand, opens the pores of the scorpion again. "
Other interesting stories were spread about the origins of the scorpions. In ancient times, according to Otto Keller's Antike Tierwelt, the idea was that scorpions would grow from dead crocodiles or (according to Pliny ) from buried sea crabs when the sun wandered through the constellation of Cancer . According to Paracelsus , scorpions emerged from rotting conspecifics, which they thereby killed. The idea that the young scorpions eat their way out of the mother's belly is similar.
In German superstition, the scorpion is said to fly at night and let everything it touches dry up. The animal already appears as the constellation Scorpio in the Babylonian calendar drawings . In fortune-telling , the scorpion meant an early death.
Scorpions in Cultural History
Scorpios have played a role in cultural history for thousands of years. They are usually portrayed in legends and myths as dangerous, deadly beings. Scorpio I is the earliest king of ancient Egypt known by name, which means that a scorpion stands at the beginning of human history, so to speak.
The Sumerian Epic of Gilgamesh already tells of scorpion people whose upper body is human and the lower body is scorpion. When the hero Gilgamesh enters Mount Mâschu, he is blocked by such a scorpion man and a scorpion woman whose "fearfulness is monstrous, the sight of whom is death". They guard the entry and exit of the sun there. No one has ever dared to go this way, but Gilgamesh manages to convince the scorpions to let him pass.
The Egyptian mythology knows the goddess Selket which is always depicted with a scorpion on her head. It has magical healing powers and protects against scorpion stings, which is why it is invoked in appropriate protective spells. When the goddess Isis is threatened by Seth , she sends seven scorpions to protect them from him. Hededet has the lower body of a scorpion. The Punic god of healing Shadrapa was also represented with a snake or a scorpion.
In Greek mythology , the angry goddess Artemis spawns a scorpion who kills the hunter Orion with one sting. Both are then moved together as constellations in the sky. Their enmity lives on there because you never see the two constellations at the same time.
Scorpions also appear in the biblical Revelation of John during the Apocalypse . The fifth trumpet calls out locusts, which receive a power "like scorpions have on earth" ( Rev 9 : 3 EU ). The pain they cause is compared to that of a stinging scorpion ( Rev 9,5 EU ), because they have tails with spines like the scorpions and use them to harm people for five months ( Rev 9,10 EU ).
"Oh, full of scorpions is my mind" says MacBeth in Shakespeare's drama (Act III, Scene 2) after the insidious dagger murder. Gottlieb Konrad Pfeffel's poem Scorpio and Shepherd Boy warns of evil people. Anna Elisabet Weirauch wrote The Scorpion (1919), Anna-Leena Härkönen (see Regine Pirschel ) The Sad Scorpion (1989). Bruna Surfistinhas The Sweet Poison of the Scorpion (2007) deals with prostitution. Hinrich Matthiesens The Scorpion (2015) is a thriller. Scorpion stings appear in John Steinbeck's novella Die Perle and the films Das Blaue Paradies (1982) or The Last Mätresse (2007). With a view to ancient Egyptian mythology or the constellation of the same name, scorpions are suitable for fantasy as well as science fiction works. In film titles, the word stands for action, paired with treachery, also in connection with sexuality: The golden scorpion (1921), the blonde scorpion (1958), the tail of the scorpion (1971), Sasori - Scorpion (1972), Scorpio, the Killer (1973), Hungry Scorpions (1985, see Julia Kent (actress) ), Red Scorpion (1989), The Hunt for the Golden Scorpion (1991), The Scorpion (1997) , Bichhoo (2000), The Black Scorpion (2000 ), Under the Spell of Jade Scorpion (2001), The Scorpion King (2002), The Sting of Scorpion (2004), Scorpion - The Fighter (2007), Scorpion (TV series, since 2014). In Rote Laterne (1991), the intriguer has “the face of a Buddha, but the heart of a scorpion.” In the action film Drive (2011) , the agile, lonely hero's jacket shows a scorpion on his back. Thrillers also cite the fable of the scorpion and the frog : The scorpion lets the frog carry him across the river and promises not to sting him, otherwise they both will die. Then he does it: So be his nature (see Wikipedia on The Scorpion and the Frog ).
Scorpion or English Scorpion is a name for motorcycles ( MZ Skorpion ), cars ( Skorpion (car brand) , Scorpion (Humberstone) , Scorpion (Innes Lee) , Ford Scorpio , Mahindra Scorpio ), fighter planes ( Sukhoi Su-25 , EADS Scorpio , Textron AirLand Scorpion ), a submachine gun , tanks ( Scorpion , Skorpion mine throwing system ), combat ships (e.g. the SMS Scorpion ) and an antique gun . Scorpions are sports clubs and rock bands, Cor Scorpii is a metal band. Scorpion- like construction of a machine is less common, such as the Claas Scorpion telehandler , the ROV Scorpio diving robotor a walking robot . Naming for space travel probably alludes to the constellation Scorpio , z. B. Richard Morgan's novel Scorpio (2007), the film Hour of the Scorpion (1968).
swell
Individual evidence
- ↑ a b c d e f Zafar Shamoon, Ryan J. Peterfy, Sami Hammoud, Babak Khazaeni: Scorpion Toxicity. StatPearls Publishing LLC, July 31, 2019, accessed June 15, 2020 .
- ^ Wilson R. Lourenço: The evolution and distribution of noxious species of scorpions (Arachnida: Scorpiones) . In: Journal of Venomous Animals and Toxins including Tropical Diseases . tape 24 , no. 1 , December 2018, ISSN 1678-9199 , p. 1 , doi : 10.1186 / s40409-017-0138-3 , PMID 29308066 , PMC 5753573 (free full text) - ( biomedcentral.com [accessed June 15, 2020]).
- ↑ Jan Ove Rein: The Scorpion Files , accessed January 19, 2018.
- ^ Scorpion poisons. In: Spektrum.de. Spektrum Akademischer Verlag, accessed June 15, 2020 .
- ^ A b c Johann J. Kleber, Ph. Philipp Wagner, Norbert Felgenhauer, Marc Kunze, Thomas Zilker: Poisoning by scorpion stings . Ed .: Deutsches Ärzteblatt. tape 96 , no. 25 , p. 1710-1715 .
- ↑ Rouhullah Dehghani, Fatemeh Kamiabi, Malihe Mohammadi: Scorpionism by Hemiscorpius spp. in Iran: a review . In: Journal of Venomous Animals and Toxins including Tropical Diseases . tape 24 , no. 1 , December 2018, ISSN 1678-9199 , p. 8 , doi : 10.1186 / s40409-018-0145-z , PMID 29507581 , PMC 5833132 (free full text) - ( biomedcentral.com [accessed June 15, 2020]).
- ↑ R. Stockmann & E. Ythier: Scorpions of the World. NAP Editions 2010, ISBN 978-2-913688-11-7 : pp. 137-141
- ↑ R. Stockmann & E. Ythier: Scorpions of the World. NAP Editions 2010, ISBN 978-2-913688-11-7 : pp. 141-144
- ↑ Isometrus deharvengi in the IUCN Red List of Endangered Species 2017.2. Posted by: L. Deharveng, A. Bedos, 2016. Retrieved September 24, 2017 ..
- ↑ Chiromachus ochropus in the IUCN Red List of Threatened Species 2017.2. Posted by: J. Gerlach, 2012. Retrieved September 24, 2017 ..
- ↑ Lychas braueri in the IUCN Red List of Threatened Species 2017.2. Posted by: J. Gerlach, 2012. Retrieved September 24, 2017 ..
- ↑ eurekalert.org of September 2, 2013: Oldest land-living animal from Godwana found.
- ^ Jan Ove Rein: The Scorpion Files. Buthidae , accessed January 19, 2018.
- ↑ R. Stockmann & E. Ythier: Scorpions of the World. NAP Editions 2010, ISBN 978-2-913688-11-7
- ↑ Victor Fet, Michael E. Soleglad, Sergei L. Zonstein: The Genus Akrav Levy, 2007 (Scorpiones: Akravidae) Revisited . In: Euscorpius. Occasional Publications in Scorpiology , Number 134, November 2011, ISSN 1536-9307 , Online PDF, 12.9 MB .
literature
Overall representations
- Arachne - the specialist magazine for arachnids . German Arachnological Society. Printing and paper Meyer, Scheinfeld 1.1995ff. ISSN 1613-2688 .
- ME Braunwalder: Scorpiones. Fauna Helvetica. Vol 13. CSCF / SZKF, Neuchâtel 2005, ISBN 2-88414-025-5 .
- J. Leeming: Scorpions of Southern Africa. Struik Publishers, Cape Town 2003, ISBN 1-86872-804-8 .
- Dieter Mahsberg, R. Lippe, S. Kallas: Scorpions. Natur und Tier-Verlag, Münster 1999, ISBN 3-931587-15-0 .
- Gary A. Polis (Ed.): The Biology of Scorpions. Stanford University Press, California 1990, ISBN 0-8047-1249-2 .
- P Weygoldt: Chelicerate, arachnids. in: Wilfried Westheide, Rieger (Ed.): Special Zoology. Part 1. Protozoa and invertebrates. Gustav Fischer Verlag, Stuttgart 1997, Spektrum Akademischer Verlag, Heidelberg 2004, ISBN 3-8274-1482-2 .
- Günter Schmidt: Scorpions and other arachnids. Landbuch-Verlag, Hannover 1992, 1996, 1999, ISBN 3-7842-1313-8 .
Special literature
- W. Bücherl: Classification, Biology and Venom Extraction of Scorpions. in: Wolfgang Bücherl, Eleanor E. Buckley: Venomous Animals and their Venoms. Academic Press, New York 1971, pp. 317-348, ISBN 0-12-138902-2 .
- V. Fet, WD Sissom, G. Lowe & ME Braunwalder: Catalog of the Scorpions of the World (1758-1998). The New York Entomological Society, New York 2000, ISBN 0-913424-24-2 .
- EN Kjellesvig-Waehring: A restudy of the fossil Scorpionida of the World. in: Palaeontographica Americana. Palaeontological Research Institution, Ithaca / New York 55.1986, pp. 1-287, ISSN 0078-8546 .
- EE Ruppert, RS Fox, RP Barnesm: Invertebrate Zoology - A functional evolutionary approach. (Chapter 18.) Brooks / Cole 2004, p. 565, ISBN 0-03-025982-7 .
- ME Soleglad, V. Fet: High-level systematics and phylogeny of the extant scorpions (Scorpiones: Orthosterni). in: Euscorpius. (electronic resource only) Marshall University, Huntington / W. Va. 11.2003, pp. 1-175 (download).
- Euscorpius (magazine)
Cultural history
- Hanns Bächtold-Stäubli (ed.): Concise dictionary of German superstition . Walter de Gruyter, Berlin 1927–42, 1987, 2000 (reprint), ISBN 3-11-016860-X .
