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Negative (very dangerous to humans) and positive (they hunt most insects, small vertebrates, which helps control pest populations for humans). (1)
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Brief Summary
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Solifuges are small spider-like arachnids that are typically associated with arid habitats with little vegetation. The approximately 1100 known solifuge species (Harvey 2002, 2003) mainly inhabit tropical and subtropical desert regions of Africa, Asia, and the Americas, but some species are also found in dry grasslands (Brookhart and Brookhart 2006). Most species are essentially crepuscular (active at dusk and dawn) or nocturnal, although some are active during the hotter parts of the day, presumably explaining the common name "sun spider". Among other common names, solifuges are also known as wind scorpions, a reference to their impressive sprinting ability, and camel spiders, a reference to the prominent arch-shaped plate on the dorsal surface of the cephalothorax of many species. Solifuges have a pair of enormous jointed pincer-like chelicerae ("jaws") which project in front of the animal between the pedipalps. The pedipalps themselves are elongate and leg-like in form. They are primarily tactile in function and are often extended in front of the body, where they are tapped on the substrate when the animal is searching for prey or (in the case of males) mates. Although they lack claws, the pedipalps have an adhesive sucker, or "palpal organ", which facilitates the grasping of prey and can even be used in climbing. Like other arachnids, solifuges have four pairs of legs, but the first pair of legs are generally reduced in size and function mainly as tactile structures. (Punzo 1998 and references therein; Brusca and Brusca 2003)

Solifuges are ferocious predators, feeding on other ground-dwelling arthropods as well as small lizards, snakes, and rodents. All or nearly all species lack venom, in lieu of which they simply tear apart their live prey with their powerful chelicerae. Some solifuges construct burrows in which they spend much of their time when not hunting or mate-searching and others hide under stones or in crevices. Females of some species guard their eggs and young offspring, which are at first gregarious but soon take on a solitary existence (sometimes after first snacking on a few siblings). (Punzo 1998 and references therein; Brusca and Brusca 2003)

Although mating has been observed in just a few species, the following likely describes the general pattern. Before copulation, the male seizes the female. In some species, he strokes and palpates her, bringing her to a passive state. He then bends her abdomen upward and opens her genital orifice with his chelicerae. Next, he produces a sperm-containing spermatophore, picks it up with his chelicerae, deposits it in her genital orifice, and leaps away. This entire sequence takes just a few minutes. In some solifuges, sperm transfer is direct, although there is still precopulatory behavior and the male inserts his chelicerae into the female orifice before and after sperm transfer. The female deposits 50 to 200 eggs in burrows in the ground or in other protected areas. (Barnes 1987) Hruskova-Martisova et al. (2010) investigated mating behavior in two sexually cannibalistic solifuges. In one species, Gluvia dorsalis, forced copulation appeared to be the only mating strategy, whereas in the other solifuge studied, Galeodes caspius subfuscus, males engaged not only in coercive mating that caused injury to their mate, but also in courtship behaviors that induce an immobile state prior to copulation (stroking with pedipalps) and during copulation (stroking and tapping). The authors suggest that coercive mating in solifuges may have evolved as an anti-predation strategy, given that sexual cannibalism occurred in around 40% of all sexual interactions observed.

Solifuges are unfortunately probably best known to the general public as the focus of alarming (and false) urban legends about them, as described here.

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Comprehensive Description
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Solifuges are small (<1 cm to 7 cm) spider-like arachnids that are typically associated with arid habitats with little vegetation. They are generally believed to be the sister group to the Pseudoscorpiones (pseudoscorpions) (Harvey 2002 and references therein). The approximately 1100 known solifuge species (Harvey 2002, 2003) mainly inhabit tropical and subtropical desert regions of Africa, Asia, and the Americas, but some species are also found in dry grasslands (Brookhart and Brookhart 2006). Most species are essentially crepuscular (active at dusk and dawn) or nocturnal, although some are active during the hotter parts of the day, presumably explaining the common name "sun spider". Among other common names, solifuges are also known as wind scorpions, a reference to their impressive sprinting ability, and camel spiders, a reference to the prominent arch-shaped plate on the dorsal surface of the cephalothorax of many species. Solifuges have a pair of enormous jointed pincer-like chelicerae ("jaws") which project in front of the animal between the pedipalps. The pedipalps themselves are elongate and leg-like in form. They are primarily tactile in function and are often extended in front of the body, where they are tapped on the substrate when the animal is searching for prey or (in the case of males) mates. Although they lack claws, the pedipalps have an adhesive sucker, or "palpal organ", which facilitates the grasping of prey and can even be used in climbing. Like other arachnids, solifuges have four pairs of legs, but the first pair of legs are generally reduced in size and function mainly as tactile structures. One of the most distinctive features of solifuges is the presence of fan-shaped stalked sensory structures (raquet organs) on the ventral surfaces of the fourth pair of legs. (Punzo 1998 and references therein; Brusca and Brusca 2003)

Solifuges are ferocious predators, feeding on other ground-dwelling arthropods as well as small lizards, snakes, and rodents. Solifuge sprinting speed has been clocked at around 50 cm/second--slower than a speedy cockroach, but faster than many potential prey. Some species are excellent climbers. All or nearly all species lack venom, in lieu of which they simply tear apart their live prey with their powerful chelicerae. Some solifuges construct burrows in which they spend much of their time when not hunting or mate-searching and others hide under stones or in crevices. Females of some species guard their eggs and young offspring, which are at first gregarious but soon take on a solitary existence (sometimes after first snacking on a few siblings). (Punzo 1998 and references therein; Brusca and Brusca 2003)

Although mating has been observed in just a few species, the following likely describes the general pattern. Before copulation, the male seizes the female. In some species, he strokes and palpates her, bringing her to a passive state. He then bends her abdomen upward and opens her genital orifice with his chelicerae. Next, he produces a sperm-containing spermatophore, picks it up with his chelicerae, deposits it in her genital orifice, and leaps away. This entire sequence takes just a few minutes. In some solifuges, sperm transfer is direct, although there is still precopulatory behavior and the male inserts his chelicerae into the female orifice before and after sperm transfer. The female deposits 50 to 200 eggs in burrows in the ground or in other protected areas. (Barnes 1987) Hruskova-Martisova et al. (2010) investigated mating behavior in two sexually cannibalistic solifuges. In one species, Gluvia dorsalis, forced copulation appeared to be the only mating strategy, whereas in the other solifuge studied, Galeodes caspius subfuscus, males engaged not only in coercive mating that caused injury to their mate, but also in courtship behaviors that induce an immobile state prior to copulation (stroking with pedipalps) and during copulation (stroking and tapping). The authors suggest that coercive mating in solifuges may have evolved as an anti-predation strategy, given that sexual cannibalism occurred in around 40% of all sexual interactions observed. Wharton (1986) reviewed the limited available literature on the mating behavior of solifuges.

Harvey (2003) brought up to date the taxonomic status and summarized the general distribution of the world Solifugae fauna and includes a dichotomous key to families. Brookhart and Brookhart (2006) reviewed the composition of the Solifugae fauna of continental North America (Canada to Mexico), recording nearly 200 species, many of which have been collected only once. Brookhart and Cushing (2008) added several species to this list with their revision of the Hemerotrecha banksi group of apparently diurnal North American solifuges.

Punzo (1998) reviewed the biology of the Solifugae. An excellent resource for anyone interested in learning more about solifuges is the Solifugae website, which provides a superb entry to the literature on this group. Solifuges are unfortunately probably best known to the general public as the focus of alarming (and false) urban legends about them, as described here.

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Solifugae
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"Camel spiders" redirects here. For the 2011 film, see Camel Spiders (film).

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 153 genera. Despite the common names, they are neither true scorpions (order Scorpiones) nor true spiders (order Araneae). Much like a spider, the body of a solifugid has two tagmata: an opisthosoma (abdomen) behind the prosoma (that is, in effect, a combined head and thorax). At the front end, the prosoma bears two chelicerae that, in most species, are conspicuously large. The chelicerae serve as jaws and in many species also are used for stridulation. Unlike scorpions, solifugids do not have a third tagma that forms a "tail". Most species of Solifugae live in dry climates and feed opportunistically on ground-dwelling arthropods and other small animals. The largest species grow to a length of 12–15 cm (5–6 in), including legs. A number of urban legends exaggerate the size and speed of the Solifugae, and their potential danger to humans, which is negligible.

Anatomy

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Ventral aspect of a solifugid, showing respiratory slots

Solifugae are moderately small to large arachnids (a few millimeters to several centimeters in body length), with the larger species reaching 12–15 cm (5–6 in) in length, including legs.[1][2] In practice, the respective lengths of the legs of various species differ drastically, so the resulting figures are often misleading. More practical measurements refer primarily to the body length, quoting leg lengths separately, if at all. The body length is up to 7 cm (3 in).[3][4] Most species are closer to 5 cm (2 in) long, and some small species are under 1 cm (0.4 in) in head-plus-body length when mature.[5]

Like that of the spider order, the Araneae, the body plan of the Solifugae has two main tagmata: the prosoma, or cephalothorax, is the anterior tagma, and the 10-segmented abdomen, or opisthosoma, is the posterior tagma. As shown in the illustrations, the solifugid prosoma and opisthosoma are not separated by nearly as clear a constriction and connecting tube or "pedicel" as occurs in "true spiders", the order Araneae. The lack of the pedicel reflects another difference between the Solifugae and spiders, namely that solifugids lack both spinnerets and silk, and do not spin webs. Spiders need considerable mobility of their abdomens in their spinning activities, and the Solifugae have no need for any such adaptation.

The prosoma comprises the head, the mouthparts, and the somites that bear the legs and the pedipalps. The alternative name "cephalothorax" reflects the fact that the prosoma includes the parts that in insects form the head plus the thorax. Though it is not split into two clear tagmata, the prosoma does have a large, relatively well-defined anterior carapace, bearing the animal's eyes and chelicerae, while a smaller posterior section bears the legs.[5][6]

Like pseudoscorpions and harvestmen, the Solifugae lack book lungs, having instead a well-developed tracheal system that inhales and exhales air through three pairs of slits on the animal's underside.

Chelicerae

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Lateral aspect of chelicera, showing teeth and cutting edge

Among the most distinctive features of the Solifugae are their large chelicerae, which in many species are longer than the prosoma. Each of the two chelicerae has two articles (segments, parts connected by a joint),[7] forming a powerful pincer, much like that of a crab; each article bears a variable number of teeth, largely depending on the species.[5][6] The chelicerae of many species are surprisingly strong; they are capable of shearing hair or feathers from vertebrate prey or carrion, and of cutting through skin and thin bones such as those of small birds.[8] Many Solifugae stridulate with their chelicerae, producing a rattling noise.[2]

Legs and pedipalps

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Male solifugid in South African veld: Its flagella are visible near the tips of the chelicerae, looking like large, backward-curling bristles. As in most species, it holds its pedipalps clear of the ground; its front legs serve as tactile sensors, barely touching the ground with their setae.

Like most other arachnids, although Solifugae appear to have five pairs of legs, only the hind four pairs actually are "true" legs. Each true leg has seven segments: coxa, trochanter, femur, patella, tibia, metatarsus, and tarsus.[8][9]

The first, or anterior, of the five pairs of leg-like appendages are not "actual" legs, but pedipalps, and they have only five segments each. The pedipalps of the Solifugae function partly as sense organs similar to insects' antennae, and partly in locomotion, feeding, and fighting. In normal locomotion, they do not quite touch the ground, but are held out to detect obstacles and prey; in that attitude, they look particularly like an extra pair of legs or perhaps arms. Reflecting the great dependence of the Solifugae on their tactile senses, their anterior true legs commonly are smaller and thinner than the posterior three pairs. That smaller anterior pair acts largely in a sensory role as a supplement to the pedipalps, and in many species they accordingly lack tarsi. At the tips of their pedipalps, Solifugae bear eversible adhesive organs, which they may use to capture flying prey, and which at least some species certainly use for climbing smooth surfaces.[8][10]

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A female of a species in the family Solpugidae showing the malleoli beneath the posterior pair of legs

For the most part, only the posterior three pairs of legs are used for running.[6][8] On the undersides of the coxae and trochanters of the last pair of legs, Solifugae have fan-shaped sensory organs called malleoli or racquet (or racket) organs. Sometimes, the blades of the malleoli are directed forward, sometimes not. They have been suspected to be sensory organs for the detection of vibrations in the soil, perhaps to detect threats and potential prey or mates.[8] These structures may be chemoreceptors.[11]

Males are usually smaller than females, with relatively longer legs.[2] Unlike females, the males bear a pair of flagella, one on each chelicera. In the accompanying photograph of a male solifugid, one flagellum is just visible near the tip of each chelicera. The flagella, which bend back over the chelicerae, are sometimes called horns and are believed to have some sexual connection, but their function has not yet been clearly explained.[8]

Eyes

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Solifugid eyes with presumably protective bristles

Some species have very large central eyes. They look like simple eyes or ocelli, and are surprisingly sophisticated. They can recognise forms, and are used in hunting and avoiding enemies. These eyes are remarkable in their internal anatomy; they may represent the last step in the integration of the aggregate of simple ocelli into a compound eye, and of further integration of a compound eye into a simple eye[clarify].[12] In contrast, lateral eyes are absent in many species, and where they are present at all, they are only rudimentary.

Classification

The Solifugae are an order of their own, though are sometimes confused with spiders, which form a completely distinct order, the Araneae. The order comprises over 1000 described species in 153 genera assigned to the following 12 families:[13]:213

The family Protosolpugidae is only known from one fossil species from the Pennsylvanian.

Ecology

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Gluvia dorsalis eating a cabbage bug (Eurydema oleracea)

Although the Solifugae are considered to be endemic indicators of desert biomes,[5]:1 they occur widely in semidesert and scrub. Some species also live in grassland or forest habitats. Solifugae generally inhabit warm and arid habitats, including virtually all warm deserts and scrublands in all continents except Antarctica and Australia.[2]

Solifugae are carnivorous or omnivorous, with most species feeding on termites, darkling beetles, and other small, ground-dwelling arthropods. Solifuges are aggressive hunters and voracious opportunistic feeders and have been recorded as feeding on snakes, small lizards, and rodents.[5] Prey is located with the pedipalps and killed and cut into pieces by the chelicerae. The prey is then liquefied and the liquid ingested through the pharynx. Although they do not normally attack humans, their chelicerae can penetrate human skin, and painful bites have been reported.[2]

Various other predators, such as the large slit-faced bat, scorpions, toads, and insectivores, may prey on Solifugae.

Life cycle

Solifugae are typically univoltine.[5]:8 Reproduction can involve direct or indirect sperm transfer; when indirect, the male emits a spermatophore on the ground and then inserts it with his chelicerae in the female's genital pore. To do this, he flings the female on her back.

The female then digs a burrow, into which she lays 50 to 200 eggs – some species then guard them until they hatch. Because the female does not feed during this time, she will try to fatten herself beforehand, and a species of 5 cm (2.0 in) has been observed to eat more than 100 flies during that time in the laboratory.[2] Solifugae undergo a number of stages including, egg, postembryo, 9–10 nymphal instars, and adults.[5]

Etymology

The name Solifugae derives from Latin, and means "those that flee from the sun". The order is also known by the names Solpugida, Solpugides, Solpugae, Galeodea, and Mycetophorae. Their common names include camel spider, wind scorpion, scorpion carrier, jerrymunglum,[14] sun scorpion, and sun spider. In southern Africa, they are known by a host of names, including red romans, haarskeerders ("hair cutters") and baardskeerders ("beard cutters"), the latter two relating to the belief they use their formidable jaws to clip hair from humans and animals to line their subterranean nests.[15]

Solifugids and humans

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A scorpion (left) fighting a solifugid (right)

Solifugids have been recognised as distinct taxa from ancient times. The Greeks recognized that they were distinct from spiders; spiders were called ἀράχνη (arachne) while Solifugae were named φαλάνγιον (phalangion). In Aelian's De natura animalium, they are mistakenly mentioned, along with scorpions, as responsible for the abandoning of a country in Ethiopia. Anton August Heinrich Lichtenstein theorised in 1797 that the "mice" that plagued the Philistines in the Old Testament were Solifugae. During World War I, troops[clarification needed] stationed in Abū Qīr, Egypt, would stage fights between captive "jerrymanders," as they referred to them, and placed bets on the outcome. Similarly, British troops stationed in Libya in World War II would stage fights between solifugids and scorpions.[5]:2–3

Urban legends

Solifugae are the subject of many legends and exaggerations about their size, speed, behaviour, appetite, and lethality. They are not especially large, the biggest having a leg span of about 12 cm (4.7 in).[2] They are fast on land compared to other invertebrates, with their top speed estimated to be 16 km/h (10 mph),[1] close to one-half as fast as the fastest human sprinter.[16]

The Solifugae apparently have neither venom glands nor any venom-delivery apparatus such as the fangs of spiders, stings of wasps, or venomous setae of caterpillars (e.g., Lonomia or Acharia species).[citation needed] One 1978 study is frequently quoted, in which the authors report detection of an exception to this rule in India, in that Rhagodes nigrocinctus had venom glands, and that injection of the secretion into mice was frequently fatal. However, no supporting studies have confirmed either statement, such as by independent detection of the glands as claimed, or the relevance of the observations, if correct. Even the authors of the original account denied having found any means of delivery of the putative venom by the animal, and the only means of administering the material to the mice was by parenteral injection.[17] Given that many non-venoms such as saliva, blood and glandular secretions can be lethal if injected, and that no venomous function was even speculated upon in this study, there is still no evidence for even one venomous species of solifugid.[18]

Because of their unfamiliar spider-like appearance and rapid movements, Solifugae have startled or even frightened many people. This fear was sufficient to drive a family from their home when one was discovered in a soldier's house in Colchester, England, and caused the family to blame the solifugid for the death of their pet dog.[19] An Arizona resident developed painful lesions due to a claimed solifugid bite, but could not produce a specimen for confirmation.[20] Though they are not venomous, the powerful chelicerae of a large specimen may inflict a painful nip, but nothing medically significant.[21]

References

  1. ^ a b "Egyptian giant solpugid (camel spider) Galeodes arabs". National Geographic. Retrieved June 10, 2011..mw-parser-output cite.citation{font-style:inherit}.mw-parser-output q{quotes:"""""'"'"}.mw-parser-output code.cs1-code{color:inherit;background:inherit;border:inherit;padding:inherit}.mw-parser-output .cs1-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Lock-green.svg/9px-Lock-green.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-limited a,.mw-parser-output .cs1-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Lock-gray-alt-2.svg/9px-Lock-gray-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Lock-red-alt-2.svg/9px-Lock-red-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration{color:#555}.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration span{border-bottom:1px dotted;cursor:help}.mw-parser-output .cs1-hidden-error{display:none;font-size:100%}.mw-parser-output .cs1-visible-error{font-size:100%}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration,.mw-parser-output .cs1-format{font-size:95%}.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-right{padding-right:0.2em}
  2. ^ a b c d e f g G. Schmidt (1993). Giftige und gefährliche Spinnentiere (in German). Westarp Wissenschaften. ISBN 3-89432-405-8.
  3. ^ Pechenik, Jan (1996). Biology of the Invertebrates. Dubuque: Wm. C. Brown Publishers. ISBN 0-697-13712-0.
  4. ^ Mullen, Gary R. (2009). Medical and Veterinary Entomology (2 ed.). Burlington, Massachusetts: Academic Press. ISBN 978-0-12-372500-4.
  5. ^ a b c d e f g h Fred Punzo (1998). The Biology of Camel-Spiders. Springer. ISBN 0-7923-8155-6. Retrieved January 25, 2010.
  6. ^ a b c Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 613–614. ISBN 0-03-056747-5.
  7. ^ Brown, Lesley (1993). The New shorter Oxford English dictionary on historical principles. Oxford: Clarendon Press. ISBN 0-19-861271-0.
  8. ^ a b c d e f Holm, Erik; Dippenaar-Schoeman, Ansie (2010). Goggo Guide: The arthropods of southern Africa. Pretoria: LAPA Publishers. ISBN 0799346896.
  9. ^ Filmer, Martin (1997). Southern African Spiders. City: BHB International / Struik. ISBN 1-86825-188-8.
  10. ^ Harmer, Sir Sidney Frederic; Shipley, Arthur Everett et alia: The Cambridge natural history Volume 4, Crustacea, Trilobites, Arachnida, Tardigrada, Pentastomida etc. Macmillan Company 1895
  11. ^ Punzo, Fred (1998). The Biology of Camel-Spiders: Arachnida, Solifugae. Boston, MA: Springer US. p. 66. ISBN 9781461557272. Retrieved 21 November 2015.
  12. ^ Beklemishev, Vladimir (1969). Principles of Comparative Anatomy of Invertebrates. Chicago: University of Chicago Press. ISBN 0226041751.
  13. ^ Levin, Simon A. (2001). Encyclopedia of biodiversity, Volume 1. 2001: Academic Press. p. 943. ISBN 978-0-12-226866-3.
  14. ^ Skaife, Sydney Harold; South African Nature Notes, Second edition. Pub: Maskew Miller: Cape Town, 1954.
  15. ^ Ross Piper (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Greenwood Press.
  16. ^ "IAAF (International Association of Athletics Federations) Biomechanical Research Project: Berlin 2009" (PDF). Archived from the original (PDF) on 2014-05-14. Retrieved 2013-11-18.
  17. ^ M. Aruchami & G. Sundara Rajulu (1978). "An investigation on the poison glands and the nature of the venom of Rhagodes nigrocinctus (Solifugae: Arachnida)". Natl. Acad. Sci. Lett. 1: 191–192.
  18. ^ Klann, Anja Elisabeth. Histology and ultrastructure of solifuges comparative studies of organ systems of solifuges (Arachnida, Solifugae) with special focus on functional analyses and phylogenetic interpretations Dissertation: Greifswald, Univ., Diss., 2009 Edition/Format:Thesis/dissertation Manuscript: eBook Archival Material: English View all editions and formats Database:WorldCat. [1]
  19. ^ "Stowaway Afghan spider kills family dog". CNN. August 28, 2008. Retrieved January 8, 2011.
  20. ^ "Mystery bug bite leaves Arizona man covered in bruises, 'excruciating pain'". Global News. July 27, 2017. Retrieved July 27, 2017.
  21. ^ David Penney (2009). "Solifugae (camel spiders)". Common Spiders and Other Arachnids of The Gambia, West Africa. Siri Scientific Press. p. 71. ISBN 978-0-9558636-3-9.

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Solifugae: Brief Summary
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"Camel spiders" redirects here. For the 2011 film, see Camel Spiders (film).

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 153 genera. Despite the common names, they are neither true scorpions (order Scorpiones) nor true spiders (order Araneae). Much like a spider, the body of a solifugid has two tagmata: an opisthosoma (abdomen) behind the prosoma (that is, in effect, a combined head and thorax). At the front end, the prosoma bears two chelicerae that, in most species, are conspicuously large. The chelicerae serve as jaws and in many species also are used for stridulation. Unlike scorpions, solifugids do not have a third tagma that forms a "tail". Most species of Solifugae live in dry climates and feed opportunistically on ground-dwelling arthropods and other small animals. The largest species grow to a length of 12–15 cm (5–6 in), including legs. A number of urban legends exaggerate the size and speed of the Solifugae, and their potential danger to humans, which is negligible.

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