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 neither true scorpions (order Scorpiones) nor true spiders (order Araneae). 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 12–15cm (5–6in), including legs. A number of urban legends exaggerate the size and speed of the Solifugae, and their potential danger to humans, which is negligible.
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 12–15cm (5–6in) in length, including legs.[2][3] 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 7cm (3in).[4][5] Most species are closer to 5cm (2in) long, and some small species are under 1cm (0.4in) in head-plus-body length when mature.[6]
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. 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.[8][6][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] As embryos they also have opisthosomal protuberances resembling the pulmonary sacs found in some palpigrades.[12]
Chelicerae
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),[13] forming a powerful pincer, much like that of a crab; each article bears a variable number of teeth, largely depending on the species.[6][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.[14] Many Solifugae stridulate with their chelicerae, producing a rattling noise.[3]
Legs and pedipalps
Male solifuge 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.
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.[14][15]
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.[16][14][17]
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][14] 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.[14] These structures may be chemoreceptors.[18]:66
Males are usually smaller than females, with relatively longer legs.[3] 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.[14]
Eyes
Solifuge eyes with bristles
Some species have very large central eyes. They look like simple eyes or ocelli, but are quite sophisticated. They can recognise forms, and are used in hunting and avoiding enemies. These eyes 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.[19] Lateral vestigial eyes can be found inserted into pits on the inner side of the cuticle. The lenses of these eyes are usually atrophied, but in some species there are both nerves and pigment cells present, and their function could be to detect motions or changes in light intensity.[20]
Classification and Phylogeny
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 1200 described species in 146 genera assigned to these 16 families:[21][22]
In 2023, the internal phylogeny of solifuges was resolved for the first time using phylogenomics.[23] The molecular phylogeny of the order showed evidence for two groups of families, which were recognized as suborders. Boreosolifugae comprises five families predominantly from the Northern Hemisphere (Eremobatidae, Gylippidae, Karschiidae, Galeodidae, and Rhagodidae). Australosolifugae comprises seven families predominantly from the Southern Hemisphere (Ammotrechidae, Ceromidae, Daesiidae, Hexisopodidae, Melanoblossidae, Mummuciidae, and Solpugidae). Daesiidae and Ammotrechidae were both recovered as non-monophyletic, due to the nested position of Melanoblossidae inside Daesiidae, and the nested position of Mummuciidae inside Ammotrechidae. A subsequent genomic study established three additional families: Dinorhaxidae (formerly within Melanoblossidae) was recovered as part of Boreosolifugae; and Lipophagidae (formerly within Gylippidae) and Namibesiidae (formerly within Daesiidae) were recovered as part of Australosolifugae.[22] This phylogeny is considered congruent with a Gondwanan origin for Australosolifugae and a Laurasian origin for Boreosolifugae.[23]
Although the Solifugae are considered to be endemic indicators of desertbiomes,[6]:1 they occur widely in semidesert and scrub. Some species also live in grassland or forest habitats. The Solifugae generally inhabit warm and arid habitats, including virtually all warm deserts and scrublands in all continents except Antarctica and Australia.[3]
Solifugae species are carnivorous or omnivorous, with most feeding on termites, darkling beetles, and other small, ground-dwelling arthropods. They are aggressive hunters and voracious opportunistic feeders, and have been recorded as feeding on snakes, small lizards, birds, and rodents.[6] 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.[3]
The Solifugae are typically univoltine (reproducing once a year).[6]: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 5cm (2.0in) has been observed to eat more than 100 flies during that time in the laboratory.[3] The Solifugae undergo a number of stages including, egg, postembryo, 9–10 nymphal instars, and adults.[6]
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,[24] 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.[25]
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).[26]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.[6]:2–3
Urban legends
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 12cm (4.7in).[3] They are fast on land compared to other invertebrates, with their top speed estimated to be 16km/h (10mph),[2] close to half as fast as the fastest human sprinter.[27]
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).[28] 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.[29] 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.[28]
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.[30] An Arizona resident developed painful lesions due to a claimed solifuge bite but could not produce a specimen for confirmation.[31] Though they are not venomous, the powerful chelicerae of a large specimen may inflict a painful nip, but nothing medically significant.[32]
Claims that Solifugae aggressively chase people are also untrue, as they are merely trying to stay in the shade/shadow provided by the human.[33]
Related Research Articles
The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids, as well as a number of extinct lineages, such as the eurypterids and chasmataspidids.
Arachnida is a class of joint-legged arthropods, in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroons.
Amblypygi is an order of arachnids also known as whip spiders or tailless whip scorpions, not to be confused with whip scorpions or vinegaroons that belong to the related order Thelyphonida. The name "amblypygid" means "blunt tail", a reference to a lack of the flagellum that is otherwise seen in whip scorpions. Amblypygids possess no silk glands or venom. They rarely bite if threatened, but can grab fingers with their pedipalps, resulting in thorn-like puncture injuries.
Schizomida, also known as sprickets or short-tailed whip-scorpions, is an order of arachnids, generally less than 5 millimetres (0.20 in) in length. The order is not yet widely studied. E. O. Wilson has identified schizomids as among the "groups of organisms that desperately need experts to work on them."
The Opiliones are an order of arachnids colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs. As of April 2017, over 6,650 species of harvestmen have been discovered worldwide, although the total number of extant species may exceed 10,000. The order Opiliones includes five suborders: Cyphophthalmi, Eupnoi, Dyspnoi, Laniatores, and Tetrophthalmi, which were named in 2014.
Uropygi is an arachnid order comprising invertebrates commonly known as whip scorpions or vinegaroons. They are often called uropygids. The name "whip scorpion" refers to their resemblance to true scorpions and possession of a whiplike tail, and "vinegaroon" refers to their ability when attacked to discharge an offensive, vinegar-smelling liquid, which contains acetic acid. The order may also be called Thelyphonida. Both names, Uropygi and Thelyphonida, may be used either in a narrow sense for the order of whip scorpions, or in a broad sense which includes the order Schizomida.
Ricinulei is a small order of arachnids. Like most arachnids, they are predatory, eating small arthropods. They occur today in west-central Africa (Ricinoides) and the Americas as far north as Texas. As of 2021, 91 extant species of ricinuleids have been described worldwide, all in the single family Ricinoididae. In older works they are sometimes referred to as Podogona. Due to their obscurity they do not have a proper common name, though in academic literature they are occasionally referred to as hooded tickspiders.
Pedipalps are the secondary pair of forward appendages among chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") and anterior to the first pair of walking legs.
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 as a type of pincers. Some chelicerae, such as those found on nearly all spiders, are hollow and contain venom glands, used to inject venom into prey or a perceived threat. Both pseudoscorpions and harvestmen have additional structures on their chelicerae that are used for grooming.
The Mesothelae are a suborder of spiders that includes a single extant family, Liphistiidae, and a number of extinct families. This suborder is thought to form the sister group to all other living spiders, and to retain ancestral characters, such as a segmented abdomen with spinnerets in the middle and two pairs of book lungs. Members of Liphistiidae are medium to large spiders with eight eyes grouped on a tubercle. They are found only in China, Japan, and southeast Asia. The oldest known Mesothelae spiders are known from the Carboniferous, over 300 million years ago.
Ammotrechidae is a family of solifuges distributed in the Americas and the Caribbean Islands. It includes 26 described genera and 95 species. Members of this family can be distinguished from members of other families by the absence of claws on tarsi of leg I, tarsal segmentation 1-2-2-(2-4), pedipalps with pairs of lateroventral spines, and by males having an immovable flagellum on the mesal face of each chelicerum. The propeltidium of the Ammotrechidae is recurved.
The anatomy of spiders includes many characteristics shared with other arachnids. These characteristics include bodies divided into two tagmata, eight jointed legs, no wings or antennae, the presence of chelicerae and pedipalps, simple eyes, and an exoskeleton, which is periodically shed.
Opiliones are an order of arachnids and share many common characteristics with other arachnids. However, several differences separate harvestmen from other arachnid orders such as spiders. The bodies of opiliones are divided into two tagmata : the abdomen (opisthosoma) and the cephalothorax (prosoma). Unlike spiders, the juncture between the abdomen and cephalothorax is often poorly defined. Harvestmen have chelicerae, pedipalps and four pairs of legs. Most harvestmen have two eyes, although there are eyeless species.
Tarantulas comprise a group of large and often hairy spiders of the family Theraphosidae. As of December 2023, 1,100 species have been identified, with 166 genera. The term "tarantula" is usually used to describe members of the family Theraphosidae, although many other members of the same infraorder (Mygalomorphae) are commonly referred to as "tarantulas" or "false tarantulas". Some of the more common species have become popular in the exotic pet trade. Many New World species kept as pets have setae known as urticating hairs that can cause irritation to the skin, and in extreme cases, cause damage to the eyes.
Spiders are air-breathing arthropods that have eight limbs, 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 diversity among all orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every land habitat. As of November 2023, 51,673 spider species in 136 families have been recorded by taxonomists. However, there has been debate among scientists about how families should be classified, with over 20 different classifications proposed since 1900.
Psalmopoeus irminia, also known as the Venezuelan suntiger, is a species of tarantula endemic to Venezuela, Guyana and Brazil. They were first described in 1994 by F. Saager.
Malleolus (plural: malleoli) is a fan-shaped chemoreceptor or racquet organ, an array of which are carried in pairs on the ventral or undersides of Solpugidae. They are the counterpart of pectines in scorpions, and modified walking limbs in the uropygids and amblypygids as well as the pedipalps in spiders and other arachnids. Most species have 5 pairs of malleoli on the ventral surface of the fourth pair of legs of both sexes, while juveniles and other species have 2-3 pairs.
In most animals the central pathways of olfactory systems are associated with glomerular neuropil and lack topographic mapping of sensory inputs. Among arthropods, the insect and crustacean olfactory (antennal) pathways are typical examples. Two orders of chelicerate arthropods, the scorpions and solpugids (Cl. Arachnida), present striking exceptions to this generalization. The major chemosensory organs of scorpions are the pectines, two ventral appendages that contact the substrate intermittently as the animal searches for food or mates. In solpugids chemosensory input is from the antennalized pedipalps and first leg pairs, and from ten fan-shaped malleoli extending ventrally to the substrate from the 4th leg pair. The pectinal and malleolar sensory systems have highly ordered arrangement of 105 to 106 primary chemoreceptors, with one (pectines) forming a two-dimensional array and the other (malleoli) assembled in a linear array. The spatial frequencies of these chemoreceptive inputs exceed 100/mm and 1000/mm, respectively, indicating a capacity for resolving structure of chemical deposits on substrates. Using several histological and axonal tracing techniques, the organization of pectinal and malleolar central projections has been resolved. The pectinal projection terminates posteriorly in the cephalothoracic mass and shows a high degree of topographic precision, perhaps to the level of individual receptors in the sensory field. This chemosensory 'map' is imposed on laminar cytoarchitecture posteriorly in the brain but merges anteriorly into glomerular substructures. The sensory projection from the malleoli shows less topographic order with fewer and larger glomeruli reminiscent of the insect olfactory system. These comparisons between arthropod taxa suggest that olfactory projections are, to varying degrees, typically glomerular but may evolve topographic and laminar organization when the stimulus field is of fixed form.
Chanbria is a genus of camel spiders. It consists of four species found in the Sonoran Desert in Mexico and the southwestern United States.
Hemiscorpius lepturus is a species of scorpion in the family Hemiscorpiidae. It is found in deserts of the Middle East, especially in southern Iraq and Iran, where it is sometimes referred to as the "Gadim scorpion".These scorpions have long, thin tails and wide bodies and grow to 8 cm (3.1 in) in males and 5.5 cm (2.2 in) in females, allowing them to live in tight rock crevices. They are fairly solitary creatures. H. lepturus has mainly been studied to discover the components and effects of its venom, which is highly lethal and is responsible for most deaths due to scorpion sting in the Iran area. H. lepturus is the only scorpion not in the family Buthidae that is potentially lethal to humans; the Buthidae family is the largest and most abundant family of scorpions, containing many highly venomous species.
1 2 3 Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp.613–614. ISBN0-03-056747-5.
↑ Shultz, Stanley; Shultz, Marguerite (2009). The Tarantula Keeper's Guide. Hauppauge, New York: Barron's. p.23. ISBN978-0-7641-3885-0.
↑ Ax, Peter (2000). "Solifugae". Multicellular Animals: The Phylogenetic System of the Metazoa. Translated by Kinsey, S. Berlin: Springer. pp.125–126. ISBN9783662103968.
↑ Harmer, Sir Sidney Frederic; Shipley, Arthur Everett et alia: The Cambridge natural history Volume 4, Crustacea, Trilobites, Arachnida, Tardigrada, Pentastomida etc. Macmillan Company 1895
↑ 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.
This page is based on this Wikipedia article Text is available under the CC BY-SA 4.0 license; additional terms may apply. Images, videos and audio are available under their respective licenses.
