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Solifugae

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Solifugae is an order of arachnids known variously as solifuges, sun spiders, camel spiders, and wind scorpions. The order includes more than 1,000 described species in about 147 genera. Despite the common names, they are neither true scorpions (order Scorpiones) nor true spiders (order Araneae). Because of this, it's less ambiguous to call them "solifuges". Most species of solifuge 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 solifuges, and their potential danger to humans, which is negligible.

Solifugae
Temporal range: Late Carboniferous–recent [1]
Solifuge from Arizona
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Chelicerata
Class: Arachnida
Order: Solifugae
Sundevall, 1833

Etymology

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The order's name is derived from the Latin "sol" meaning "sun" and "fugere" meaning "to flee". Put together, it means "those who flee from the sun".[2] These animals have a number of common names including sun spiders, wind scorpions, wind spiders, red romans, and camel spiders.[2] In Afrikaans, they are known as "haarskeerders" ("hair cutters"), and "baardskeerders" ("beard cutters"). This is in reference to myths that they cut hair to be used as nest bedding.[2]

Anatomy and physiology

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Ventral aspect of a solifuge, showing respiratory slots

Solifuges 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.[3][4] In practice, the respective lengths of the legs of various species differ greatly, 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).[5][6] 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.[7]

Like that of spiders, 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. The abdominal tergites and sternites are separated by large areas of intersegmental membranes, giving it a high degree of flexibility and ability to stretch considerably, which allows it to consume a large amount of food.[7] As shown in the illustrations, the solifuge prosoma and opisthosoma are not separated by nearly as clear a constriction and connecting tube or "pedicel" as occurs in Araneae. The lack of the pedicel reflects another difference between the Solifugae and spiders, namely that solifuges 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 such adaptation.

The prosoma comprises the head, the mouthparts, and the somites that bear the legs and the pedipalps. It is covered by a carapace, also called a prosomal dorsal shield or peltidium, which is composed of three distinct elements called propeltidium, mesopeltidium and metapeltidium. The propeltidium contains the eyes, the chelicerae that, in most species, are conspicuously large, the pedipalps and the first two pairs of legs. Meso- and metapeltidium contains the third and fourth pairs of legs.[7][8][9] The chelicerae serve as jaws and in many species also are used for stridulation. Unlike scorpions, solifuges do not have a third tagma that forms a "tail".

Currently, neither fossil nor embryological evidence shows that arachnids ever had a separate thorax-like division, so the validity of the term cephalothorax, which means a fused cephalon, or head, and thorax, has been questioned. Also, arguments exist against use of "abdomen", as the opisthosoma of many arachnids contains organs atypical of an abdomen, such as a heart and respiratory organs.[10]

Like other arachnids outside the orders of scorpions and the Tetrapulmonata, the Solifugae lack book lungs, having instead a well-developed tracheal system that inhales and exhales air through a number of spiracles - one pair between the second and third pair of walking legs, two pairs on the abdomen on abdominal segments three and four, and an unpaired spiracle on the fifth abdominal segment.[11] Air sacs are attached to the branching tracheae, with tracheoles penetrating the epithelia of internal organs. Hemocyanin, a respiratory pigment common in the hemolymph of many arachnids and other arthropods, is absent.[12] As embryos they also have opisthosomal protuberances resembling the pulmonary sacs found in some palpigrades.[13]

Chelicerae

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Chewing movement of the chelicerae in solifuges dorsal view (top) and side view (bottom)
 
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),[14] forming a powerful pincer, much like that of a crab; each article bears a variable number of teeth, largely depending on the species.[7][9] 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.[15] Many Solifugae stridulate with their chelicerae, producing a rattling noise.[4]

Legs and pedipalps

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Male solifuge in South African veld: Its flagella are visible near the tips of the chelicerae, looking like large, backward-curling bristles.

These elements work the same way as in most other arachnids. Although the Solifugae appear to have five pairs of legs, only the hind four pairs are true legs. Each true leg has seven segments: coxa, trochanter, femur, patella, tibia, metatarsus, and tarsus.[15][16]

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 a membranous suctorial organ, which are used for capturing prey, and also for bringing water to their mouthparts for drinking and climbing smooth surfaces.[17][15][18]

 
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.[9][15] On the undersides of the coxae and trochanters of the last pair of legs, the 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.[15] These structures may be chemoreceptors.[7]: 66 

Males are usually smaller than females, with relatively longer legs.[4] Unlike females, the males bear a pair of flagella, one on each chelicera. In the accompanying photograph of a male solifuge, 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.[15]

Eyes

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Solifuge eyes with bristly setae

Solifuges have a pair of large central eyes known as median ocelli[7]: 68  These eyes are oriented at the very front of its cephalothorax and are placed close together.[19]: 68  These eyes have a pigment-cup structure and are covered by a domed outer lens made from the animal's exoskeleton.[7]: 68–69  Below the dome is the animal's retina, a multi-tiered structure with a layer of cells called the vitreous body at its top.[7]: 68  Underneath is the thin preretinal membrane, acting as a barrier between the vitreous body above and the rhabdomeres beneath. Rhabdomeres are light-sensitive and function as the eye's photoreceptors.[7]: 68  Interspersed between the rhabdomeres are pigment cells.[7]: 68  The eye's optic nerve begins at its center and is connected to the axons of numerous rhabdomeres.[7]: 68 

In addition to the median eyes, solifuges possess a pair of vestigial lateral ocelli. These eyes are found in pits on the animal's cephalic lobes near the chelicerae.[7]: 68  The ocelli's lenses are usually atrophied. However, in some species both nerves and pigment cells are present.[7]: 68  In species where lateral eyes are functional, they probably aid in detecting motions or changes in light intensity.[7]: 68 

Habitat and distribution

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Most solifuges live in tropics and subtropical deserts in the Americas, Southern Europe, Africa, the Middle East, and South Asia. Surprisingly, these animals are absent in Australia and Madagascar.[7]: 108  Within the desert, solifuges live in a variety of micro-habitats. These include sand dunes, sand flats, floodplains, rocky hillsides, desert shrublands, gravel plains, and mountain valleys.[7]: 108  In addition to the desert, certain solifuges live in more arid grasslands and forests.[7]: 113–115 

Depending on the species in question, solifuges may be more sedentary or on the move. Sedentary species are often fossorial, living in relatively permanent burrows underground. Transitory species spend most of their time up the surface, occasionally seeking refuge in cracks or under rocks and vegetation.[7]: 108 

Behavior and life history

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Diet and hunting

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Gluvia dorsalis eating a cabbage bug (Eurydema oleracea)

Solifuges are carnivores and typically generalists, feeding on a wide variety of prey in their given environment.[7]: 131–132  For most species, insects make up the bulk of their diet.[7]: 132  However, these animals have been known to consume anything they can subdue.[7]: 132  This includes other arachnids like spiders, scorpions, and smaller solifuges, other arthropods like millipedes, and small lizards, birds, and mammals.[7]: 132  Additionally, solifuges are voracious eaters. It's common for adult females to eat so much that they're temporarily unable to walk.[7]: 132 

When looking for prey, most solifuges rapidly move about while tapping their pedipalps on the ground.[7]: 151  The only exception is the majority of termite-loving species, as they prefer to be more sedentary.[7]: 151  In addition to using their pedipalps, solifuges have a variety of methods to locate prey. These include seeing movements with their eyes, feeling with their long hairlike setae, smelling with their malleoli, and sensing vibrations.[7]: 151–152  How much the animal relies on each sense depends on the species.[7]: 152  While all hunt on the ground, some species are great climbers, able to search for prey on trees, shrubs, and on artificial structures.[7]: 156 

Solifuges hunt their prey using three main hunting strategies: stalking, chasing, and ambushing.[7]: 156  Depending on the meal's size, prey is seized with the animal's pedipalps or massive chelicerae.[7]: 157  When the pedipalps are used, prey is initially caught with the limb's suction cups, then rapidly pulled towards the chelicerae to be chewed.[7]: 158  These motions happen so fast that they can't be distinguished.[7]: 158  Before eating, solifuges prepare their food by removing any parts they find unfavorable. In arthropods, these are typically areas that have a high amount of chitin (heads, antennae, wings, etc).[7]: 164 

Solifuges eat in different ways based on the shape of their food. Prey that is long and narrow is held perpendicular to the chelicerae and chewed from one end to another. More round prey is chewed by rotating the body all at once.[7]: 158  This chewing motion turns the food into a liquidized paste which is then swallowed by the animal's pharynx.[7]: 158  Solifuges that haven't fed for long periods are known to eat faster than ones that fed recently.[7]: 161  Larger solifuges are also known to eat faster than smaller ones.[7]: 161 

Reproduction

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The Solifugae are typically univoltine (reproducing once a year).[20]: 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 tries 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.[4] The Solifugae undergo a number of stages including, egg, postembryo, 9–10 nymphal instars, and adults.[20]

Classification and phylogeny

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Solifuges are an order of arachnids comprising over 1200 species in 146 genera assigned to 16 different families.[21][22] Solifuges can be divided into two groups of families which are recognized as distinct suborders.[23] These are the Australosolifugae which live predominantly in the Southern Hemisphere and the Boreosolifugae which live mostly in the Northern Hemisphere.[23] This phylogeny is considered congruent with a Gondwanan origin for Australosolifugae and a Laurasian origin for Boreosolifugae.[23] When looking at their relationships, the families Ammotrechidae and Daesiidae were found to be paraphyletic, leading to multiple clades without a name.[23] Because of this, a later genomic study established three additional families: Dinorhaxidae, Lipophagidae, and Namibesiidae.[22]

Suborder Boreosolifugae
Suborder Australosolifugae
incertae sedis

Phylogeny

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Below is a family tree of the various solifuge families based on phylogenomics.[21][22]

 Solifugae 

Relationship with humans

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A scorpion (left) fighting a solifuge (right)

Solifuges have been recognized as distinct taxa from ancient times. In Aelian's De natura animalium, "four-jawed spiders" are credited, along with scorpions, as being responsible for the abandoning of a desert region near the Astaboras river (said to be in India, but thought to be a river in Ethiopia).[24] Anton August Heinrich Lichtenstein theorized in 1797 that the "mice" that plagued the Philistines in the Old Testament were Solifugae.[citation needed] 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 staged fights between solifuges and scorpions.[7]: 2–3 

Urban legends

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The Solifugae are the subject of many legends and exaggerations about their size, speed, behavior, appetite, and lethality. They are not especially large, the biggest having a leg span around 12 cm (4.7 in).[4] They are fast on land compared to other invertebrates, with their top speed estimated to be 16 km/h (10 mph).[3]

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).[25] One 1978 study is frequently quoted, in which the authors report detection of an exception 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 admitted to having found no 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.[26] 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 solifuge.[25]

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 allegedly discovered in a soldier's house in Colchester, England, and caused the family to blame the solifuge for the death of their pet dog.[27] An Arizona resident developed painful lesions due to a claimed solifuge bite but could not produce a specimen for confirmation.[28] Though they are not venomous, the powerful chelicerae of a large specimen may inflict a painful nip, but nothing medically significant.[29]

Claims that Solifugae aggressively chase people are also untrue, as they are merely trying to stay in the shade/shadow provided by the human.[30]

References

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  1. ^ "Solifugae (camel spider)". Paleobiology Database.
  2. ^ a b c Larson, Norman. "Solifugae (solifuges, solifugids, solpugids)". www.biodiversityexplorer.info. Retrieved 1 September 2024.
  3. ^ a b "Egyptian giant solpugid (camel spider) Galeodes arabs". National Geographic. 11 April 2010. Archived from the original on January 21, 2010. Retrieved June 10, 2011.
  4. ^ a b c d e Schimdt, G. (1993). Giftige und gefährliche Spinnentiere (in German). Westarp Wissenschaften. ISBN 3-89432-405-8.
  5. ^ Pechenik, Jan (1996). Biology of the Invertebrates. Dubuque: Wm. C. Brown Publishers. ISBN 0-697-13712-0.
  6. ^ Mullen, Gary R. (2009). Medical and Veterinary Entomology (2 ed.). Burlington, Massachusetts: Academic Press. ISBN 978-0-12-372500-4.
  7. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al Fred Punzo (1998). The Biology of Camel-Spiders. Springer. ISBN 0-7923-8155-6. Retrieved 31 August 2024.
  8. ^ Dunlop, Jason A.; Krüger, Jessica; Alberti, Gerd (31 July 2012). "The sejugal furrow in camel spiders and acariform mites". Arachnologische Mitteilungen. 43: 29–36. doi:10.5431/aramit4303.
  9. ^ a b c Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 613–614. ISBN 0-03-056747-5.
  10. ^ Shultz, Stanley; Shultz, Marguerite (2009). The Tarantula Keeper's Guide. Hauppauge, New York: Barron's. p. 23. ISBN 978-0-7641-3885-0.
  11. ^ Ax, Peter (2000). "Solifugae". Multicellular Animals: The Phylogenetic System of the Metazoa. Translated by Kinsey, S. Berlin: Springer. pp. 125–126. ISBN 9783662103968.
  12. ^ Evolution of Air Breathing: Oxygen Homeostasis and the Transitions from Water to Land and Sky
  13. ^ Ballesteros, Jesús A.; Santibáñez López, Carlos E.; Kováč, Ľubomír; Gavish-Regev, Efrat; Sharma, Prashant P. (18 December 2019). "Ordered phylogenomic subsampling enables diagnosis of systematic errors in the placement of the enigmatic arachnid order Palpigradi". Proceedings of the Royal Society B: Biological Sciences. 286 (1917): 20192426. doi:10.1098/rspb.2019.2426. PMC 6939912. PMID 31847768.
  14. ^ Brown, Lesley (1993). The New shorter Oxford English dictionary on historical principles. Oxford: Clarendon Press. ISBN 0-19-861271-0.
  15. ^ a b c d e f Holm, Erik; Dippenaar-Schoeman, Ansie (2010). Goggo Guide: The arthropods of southern Africa. Pretoria: LAPA Publishers. ISBN 978-0799346893.
  16. ^ Filmer, Martin (1997). Southern African Spiders. City: BHB International / Struik. ISBN 1-86825-188-8.
  17. ^ A sticky situation: Solifugids (Arachnida, Solifugae) use adhesive organs on their pedipalps for prey capture
  18. ^ Harmer, Sir Sidney Frederic; Shipley, Arthur Everett et alia: The Cambridge natural history Volume 4, Crustacea, Trilobites, Arachnida, Tardigrada, Pentastomida etc. Macmillan Company 1895
  19. ^ Ruppert, Edward E.; Fox, Richard S.; Barnes, Robert D. (2004). Invertebrate zoology : a functional evolutionary approach. Internet Archive. Belmont, CA : Thomson-Brooks/Cole. ISBN 978-0-03-025982-1.
  20. ^ a b Fred Punzo (1998). The Biology of Camel-Spiders. Springer. ISBN 0-7923-8155-6. Retrieved January 25, 2010.
  21. ^ a b "World Solifugae Catalog". World Solifugae Catalog. Natural History Museum Bern. Retrieved 5 June 2022.
  22. ^ a b c Kulkarni, Siddharth S.; Yamasaki, Takeshi; Thi Hong Phung, Luong; Karuaera, Nanguei; Daniels, Savel R.; Gavish-Regev, Efrat; Sharma, Prashant P. (1 February 2024). "Phylogenomic data reveal three new families of poorly studied Solifugae (camel spiders)". Molecular Phylogenetics and Evolution. 191: 107989. doi:10.1016/j.ympev.2023.107989. ISSN 1055-7903. PMID 38072141. S2CID 266166984.
  23. ^ a b c d Kulkarni, Siddharth S.; Steiner, Hugh G.; Garcia, Erika L.; Iuri, Hernán; Jones, R. Ryan; Ballesteros, Jesús A.; Gainett, Guilherme; Graham, Matthew R.; Harms, Danilo; Lyle, Robin; Ojanguren-Affilastro, Andrés A.; Santibañez-López, Carlos E.; Silva de Miranda, Gustavo; Cushing, Paula E.; Gavish-Regev, Efrat (15 September 2023). "Neglected no longer: Phylogenomic resolution of higher-level relationships in Solifugae". iScience. 26 (9): 107684. Bibcode:2023iSci...26j7684K. doi:10.1016/j.isci.2023.107684. ISSN 2589-0042. PMC 10484990. PMID 37694155.
  24. ^ "Aelian, NA. 17.40". attalus.org. Retrieved March 6, 2019. - where they are called "four-jawed spiders".
  25. ^ a b Klann, Anja Elisabeth (2009). Histology and Ultrastructure of Solifuges: Comparative Studies of Organ Systems of Solifuges (Arachnida, Solifugae) with Special Focus on Functional Analyses and Phylogenetic Interpretations (Dr. rer. nat. thesis). Universität Greifswald.
  26. ^ Aruchami, M.; Sundara Rajulu, G. (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.
  27. ^ Siddique, Haroon (2008-08-28). "Paratrooper's family flees home after Afghan spider 'kills dog'". The Guardian. Retrieved 2019-08-01.
  28. ^ Sung, Jenny (July 27, 2017). "Mystery bug bite leaves Arizona man covered in bruises, 'excruciating pain'". Global News. Archived from the original on 2022-08-04. Retrieved July 27, 2017.
  29. ^ 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.
  30. ^ Szalay, Jessie (17 December 2014). "Camel Spiders: Facts & Myths". livescience.com. Purch Group. Live Science. Retrieved 19 July 2023.

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