49-596: Thelyphonida Latreille, 1804 (as Thélyphone) Uropygi is an arachnid order comprising invertebrates commonly known as whip scorpions or vinegaroons (also spelled vinegarroons and vinegarones ). 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
98-461: A bundle from a small chamber connected to the spiracle . This type of tracheal system has almost certainly evolved from the book lungs, and indicates that the tracheae of arachnids are not homologous with those of insects. Further adaptations to terrestrial life are appendages modified for more efficient locomotion on land, internal fertilisation, special sensory organs, and water conservation enhanced by efficient excretory structures as well as
147-444: A clade called Arachnopulmonata was also well supported. Pseudoscorpiones may also belong here, as all six orders share the same ancient whole genome duplication , and analyses support pseudoscorpions as the sister group of scorpions. Genetic analysis has not yet been done for Ricinulei, Palpigradi, or Solifugae, but horseshoe crabs have gone through two whole genome duplications, which gives them five Hox clusters with 34 Hox genes ,
196-470: A fourth pair usually appears when they moult into nymphs . However, mites are variable: as well as eight, there are adult mites with six or, like in Eriophyoidea , even four legs. While the adult males in some members of Podapolipidae have six legs, the adult females have only a single pair. Arachnids are further distinguished from insects by the fact they do not have antennae or wings . Their body
245-404: A narrow sense for the order of whip scorpions, or in a broad sense which includes the order Schizomida . Carl Linnaeus first described a whip scorpion in 1758, although he did not distinguish it from what are now regarded as different kinds of arachnid, calling it Phalangium caudatum . Phalangium is now used as a name for a genus of harvestmen ( Opiliones ). In 1802, Pierre André Latreille
294-508: A single order of arachnids, Pedipalpi. Pedipalpi was later divided into two orders, Amblypygi and Uropygi (or Uropygida). Schizomida was then split off from Uropygi into a separate order. The remainder has either continued to be called by the same name, Uropygi, possibly distinguished as Uropygi sensu stricto , or called Thelyphonida. When the name Uropygi is used for the whip scorpions, the clade containing Uropygi and Schizomida may be called Thelyphonida, or Thelyphonida s.l. Conversely, when
343-413: A thin membrane. Inside the pit, a small hair touches the underside of the membrane, and detects its motion. Slit sense organs are believed to be involved in proprioception , and possibly also hearing. Arachnids may have one or two gonads , which are located in the abdomen. The genital opening is usually located on the underside of the second abdominal segment. In most species, the male transfers sperm to
392-427: A transverse fold of the ectoderm . The ancestors of modern arachnids probably had both types, but modern ones often lack one type or the other. The cornea of the eye also acts as a lens, and is continuous with the cuticle of the body. Beneath this is a transparent vitreous body, and then the retina and, if present, the tapetum. In most arachnids, the retina probably does not have enough light sensitive cells to allow
441-435: A waxy layer covering the cuticle. The excretory glands of arachnids include up to four pairs of coxal glands along the side of the prosoma, and one or two pairs of Malpighian tubules , emptying into the gut. Many arachnids have only one or the other type of excretory gland, although several do have both. The primary nitrogenous waste product in arachnids is guanine . Arachnid blood is variable in composition, depending on
490-427: Is crushed between special teeth on the inside of the trochanters (the second segment of the "legs") of the front appendages. They are valuable in controlling the population of cockroaches and crickets. Males secrete a spermatophore (a united mass of sperm), which is transferred to the female following courtship behaviour, in which the male holds the ends of the female's first legs in his chelicerae . The spermatophore
539-406: Is deposited on the ground and picked up by the female using her genital area. In some genera, the male then uses his pedipalps to push the spermatophore into her body. After a few months, the female will dig a large burrow and seal herself inside. Up to 40 eggs are extruded , within a membranous broodsac that preserves moisture and remains attached to the genital operculum and the fifth segment of
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#1733085496294588-505: Is derived from the Greek word ἀράχνη ( aráchnē , 'spider'), from the myth of the hubristic human weaver Arachne , who was turned into a spider. Almost all adult arachnids have eight legs, unlike adult insects which all have six legs. However, arachnids also have two further pairs of appendages that have become adapted for feeding, defense, and sensory perception. The first pair, the chelicerae , serve in feeding and defense. The next pair,
637-403: Is organized into two tagmata , called the prosoma and opisthosoma , also referred to as the cephalothorax and abdomen . However, there are questions about the validity of the latter terms. While the term cephalothorax implies a fused cephalon (head) and thorax , there is currently neither fossil nor embryological evidence that arachnids ever had a separate thorax-like division. Likewise,
686-404: Is the posterior part of the body in some arthropods , behind the prosoma ( cephalothorax ). It is a distinctive feature of the subphylum Chelicerata ( arachnids , horseshoe crabs and others). Although it is similar in most respects to an abdomen (and is often referred to as such), the opisthosoma is differentiated by its inclusion of the respiratory organs ( book lungs or book gills ) and
735-507: The cephalothorax . In some species the frontmost pair of legs has converted to a sensory function, while in others, different appendages can grow large enough to take on the appearance of extra pairs of legs. Almost all extant arachnids are terrestrial , living mainly on land. However, some inhabit freshwater environments and, with the exception of the pelagic zone , marine environments as well. They comprise over 110,000 named species , of which 51,000 are species of spiders. The term
784-493: The Lower Cretaceous Crato Formation could be the same size. Because of their legs, claws, and "whip", though, they can appear much larger, and the heaviest specimen weighed was 12.4 grams (0.44 oz). The opisthosoma consists of 12 segments. The first segment forms a pedicel, and each of the next eight segments has dorsal tergites . The last three segments are fused into closed rings that ends with
833-509: The cladogram below. Including fossil taxa does not fundamentally alter this view, although it introduces some additional basal groups. Chelicerata (sea spiders, horseshoe crabs and arachnids ) [REDACTED] [REDACTED] [REDACTED] Myriapoda (centipedes, millipedes, and allies) [REDACTED] [REDACTED] Pancrustacea (crustaceans and hexapods) [REDACTED] [REDACTED] The extant chelicerates comprise two marine groups: Sea spiders and horseshoe crabs, and
882-484: The heart . The number of segments and appendages on the opisthosoma vary. Scorpions have 13, but the first is only seen during its embryological development. Other arachnids have fewer; harvestmen , for instance, have only ten. In general, appendages are absent or reduced, although in horseshoe crabs they persist as large plate-like limbs, called opercula or branchiophores, bearing the book gills , and that function in locomotion and gas exchange. In most chelicerates
931-424: The house dust mite , are also the only arachnids able to ingest solid food, which exposes them to internal parasites, although it is not unusual for spiders to eat their own silk. And one species of spider is mostly herbivorous. Scorpions, spiders and pseudoscorpions secrete venom from specialized glands to kill prey or defend themselves. Their venom also contains pre-digestive enzymes that helps breaking down
980-453: The pedipalps , have been adapted for feeding, locomotion, and/or reproductive functions. In scorpions, pseudoscorpions, and ricinuleids the pedipalps end in a pair of pinchers, while in whip scorpions, Schizomida , Amblypygi , and most harvestmen, they are raptorial and used for prey capture. In Solifugae , the palps are quite leg-like, so that these animals appear to have ten legs. The larvae of mites and Ricinulei have only six legs;
1029-453: The 'abdomen' of many arachnids contains organs atypical of an abdomen, such as a heart and respiratory organs. The cephalothorax is usually covered by a single, unsegmented carapace. The abdomen is segmented in the more primitive forms, but varying degrees of fusion between the segments occur in many groups. It is typically divided into a preabdomen and postabdomen, although this is only clearly visible in scorpions, and in some orders, such as
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#17330854962941078-524: The Gambia and Guinea-Bissau . They usually dig burrows with their pedipalps, to which they transport their prey. They may also burrow under logs, rotting wood, rocks, and other natural debris. They prefer humid, dark places and avoid light. Mastigoproctus giganteus , the giant whip scorpion, is found in more arid areas, including Arizona and New Mexico. As of 2023, the World Uropygi Catalog accepted
1127-436: The abdomen has no appendages. Like all arthropods, arachnids have an exoskeleton , and they also have an internal structure of cartilage -like tissue, called the endosternite , to which certain muscle groups are attached. The endosternite is even calcified in some Opiliones . Most arachnids lack extensor muscles in the distal joints of their appendages. Spiders and whip scorpions extend their limbs hydraulically using
1176-409: The arachnids has proven difficult as of March 2016 , with successive studies producing different results. A study in 2014, based on the largest set of molecular data to date, concluded that there were systematic conflicts in the phylogenetic information, particularly affecting the orders Acariformes , Parasitiformes and Pseudoscorpiones , which have had much faster evolutionary rates. Analyses of
1225-422: The burrow. The mother may live up to two more years. The young grow slowly, going through four moults in about four years before reaching adulthood. They live for up to another four years. Whip scorpions are found in tropical and subtropical areas, excluding Europe and Australia . Also, only a single species is known from Africa : Etienneus africanus , probably a Gondwana relict , endemic to Senegal ,
1274-920: The data using sets of genes with different evolutionary rates produced mutually incompatible phylogenetic trees . The authors favoured relationships shown by more slowly evolving genes, which demonstrated the monophyly of Chelicerata, Euchelicerata and Arachnida, as well as of some clades within the arachnids. The diagram below summarizes their conclusions, based largely on the 200 most slowly evolving genes; dashed lines represent uncertain placements. Acariformes [REDACTED] Opiliones [REDACTED] Ricinulei [REDACTED] Solifugae [REDACTED] Parasitiformes [REDACTED] Pseudoscorpiones [REDACTED] Scorpiones [REDACTED] Araneae [REDACTED] Amblypygi [REDACTED] Uropygi (Thelyphonida s.s. ) [REDACTED] Tetrapulmonata , here consisting of Araneae , Amblypygi and Uropygi (Thelyphonida s.s. ) ( Schizomida
1323-419: The eyes to form a proper image. In addition to the eyes, almost all arachnids have two other types of sensory organs. The most important to most arachnids are the fine sensory hairs that cover the body and give the animal its sense of touch. These can be relatively simple, but many arachnids also possess more complex structures, called trichobothria . Finally, slit sense organs are slit-like pits covered with
1372-554: The female in a package, or spermatophore . The males in harvestmen and some mites have a penis. Complex courtship rituals have evolved in many arachnids to ensure the safe delivery of the sperm to the female. Members of many orders exhibit sexual dimorphism. Arachnids usually lay yolky eggs , which hatch into immatures that resemble adults. Scorpions, however, are either ovoviviparous or viviparous , depending on species, and bear live young. Also some mites are ovoviviparous and viviparous, even if most lay eggs. In most arachnids only
1421-640: The females provide parental care, with harvestmen being one of the few exceptions. The phylogenetic relationships among the main subdivisions of arthropods have been the subject of considerable research and dispute for many years. A consensus emerged from about 2010 onwards, based on both morphological and molecular evidence; extant (living) arthropods are a monophyletic group and are divided into three main clades: chelicerates (including arachnids), pancrustaceans (the paraphyletic crustaceans plus insects and their allies), and myriapods (centipedes, millipedes and allies). The three groups are related as shown in
1470-456: The flagellum, made up of 30-40 units. Like the related orders Schizomida and Amblypygi, whip scorpions use only six legs for walking, with the first two legs serving as antennae-like sensory organs. All species also have very large scorpion-like pedipalps (pincers) but there is an additional large spine on each palpal tibia. They have one pair of median eyes at the front of the cephalothorax and up till five pairs of lateral eyes on each side of
1519-554: The following 16 extant genera, all placed in the family Thelyphonidae: In addition, seven extinct genera were accepted, two within the family Thelyphonidae: and five unplaced as to family: Arachnid Arachnids are arthropods in the class Arachnida ( / ə ˈ r æ k n ɪ d ə / ) of the subphylum Chelicerata . Arachnida includes, among others, spiders , scorpions , ticks , mites , pseudoscorpions , harvestmen , camel spiders , whip spiders and vinegaroons . Adult arachnids have eight legs attached to
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1568-489: The food. It extends through most of the body, and connects to a short sclerotised intestine and anus in the hind part of the abdomen. Arachnids have two kinds of eyes: the lateral and median ocelli . The lateral ocelli evolved from compound eyes and may have a tapetum , which enhances the ability to collect light. With the exception of scorpions, which can have up to five pairs of lateral ocelli, there are never more than three pairs present. The median ocelli develop from
1617-556: The head, a pattern also found in scorpions. Whip scorpions have no venom glands, but they have glands near the rear of their abdomen that can spray a combination of acetic acid and caprylic acid when they are bothered. The acetic acid gives this spray a vinegar-like smell, giving rise to the common name vinegaroon . Whip scorpions are carnivorous , nocturnal hunters feeding mostly on insects , millipedes , scorpions, and terrestrial isopods, but sometimes on worms and slugs. Mastigoproctus sometimes preys on small vertebrates . The prey
1666-555: The highest number found in any invertebrate, yet it is not clear if the oldest genome duplication is related to the one in Arachnopulmonata. Onychophora [REDACTED] Mandibulata [REDACTED] [REDACTED] [REDACTED] Pycnogonida [REDACTED] † Chasmataspidida [REDACTED] † Eurypterida [REDACTED] Parasitiformes [REDACTED] Acariformes [REDACTED] Pseudoscorpiones [REDACTED] Opisthosoma The opisthosoma
1715-449: The late Carboniferous , somewhere in the tropics of Pangaea . whip spiders [REDACTED] whip scorpions [REDACTED] short-tailed whip scorpions [REDACTED] Whip scorpions range from 25 to 85 mm (1.0 to 3.3 in) in length, with most species having a body no longer than 30 mm (1.2 in); the largest species, of the genus Mastigoproctus , can reach 85 mm (3.3 in). An extinct Mesoproctus from
1764-475: The mites, the abdominal sections are completely fused. A telson is present in scorpions, where it has been modified to a stinger, and into a flagellum in the Palpigradi , Schizomida (very short) and whip scorpions . At the base of the flagellum in the two latter groups there are glands which produce acetic acid as a chemical defense. Except for a pair of pectines in scorpions, and the spinnerets in spiders,
1813-486: The mode of respiration. Arachnids with an efficient tracheal system do not need to transport oxygen in the blood, and may have a reduced circulatory system. In scorpions and some spiders, however, the blood contains haemocyanin , a copper-based pigment with a similar function to haemoglobin in vertebrates. The heart is located in the forward part of the abdomen, and may or may not be segmented. Some mites have no heart at all. Arachnids are mostly carnivorous , feeding on
1862-475: The mother's ventral opisthosoma . The female refuses to eat and holds her opisthosoma in an upward arch so that the broodsac does not touch the ground for the next few months, as the eggs develop into postembryos . Appendages become visible. The white young that hatch from the postembryos climb onto their mother's back and attach themselves there with special suckers. After the first moult , when they look like miniature adults but with bright red palps, they leave
1911-416: The mouth. Behind the mouth is a muscular, sclerotised pharynx , which acts as a pump, sucking the food through the mouth and on into the oesophagus and stomach . In some arachnids, the oesophagus also acts as an additional pump. The stomach is tubular in shape, with multiple diverticula extending throughout the body. The stomach and its diverticula both produce digestive enzymes and absorb nutrients from
1960-446: The name Thelyphonida is used for the whip scorpions alone, the parent clade may be called Uropygi, or Uropygi sensu lato . The table below summarizes the two usages. When the qualifications s.l. and s.s. are omitted, the names Uropygi and Thelyphonida are ambiguous. Phylogenetic studies show the three groups, Amblypygi, Uropygi s.s. and Schizomida, to be closely related. The Uropygi s.s. and Schizomida likely diverged in
2009-438: The opisthosomal limbs are greatly reduced and persist only as specialized structures, such as the silk-producing spinnerets of spiders or the pectines of scorpions. In animals like whip scorpions and whip spiders the first two 'sternites' bearing the book lungs may actually be highly modified opisthosomal limbs. Segmentation of the opisthosoma in adult spiders is not visible, but embryo spiders typically have 13 segments,
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2058-457: The pre-digested bodies of insects and other small animals. But ticks, and many mites, are parasites, some of which are carriers of disease. The diet of mites also include tiny animals, fungi, plant juices and decomposing matter. Almost as varied is the diet of harvestmen , where we will find predators, decomposers and omnivores feeding on decaying plant and animal matter, droppings, animals and mushrooms. The harvestmen and some mites, such as
2107-460: The pressure of their hemolymph . Solifuges and some harvestmen extend their knees by the use of highly elastic thickenings in the joint cuticle. Scorpions, pseudoscorpions and some harvestmen have evolved muscles that extend two leg joints (the femur-patella and patella-tibia joints) at once. The equivalent joints of the pedipalps of scorpions though, are extended by elastic recoil. There are characteristics that are particularly important for
2156-407: The prey. The saliva of ticks contains anticoagulants and anticomplements, and several species produce a neurotoxin . Arachnids produce digestive enzymes in their stomachs, and use their pedipalps and chelicerae to pour them over their dead prey. The digestive juices rapidly turn the prey into a broth of nutrients, which the arachnid sucks into a pre-buccal cavity located immediately in front of
2205-487: The terrestrial arachnids. These have been thought to be related as shown below. (Pycnogonida (sea spiders) may be excluded from the chelicerates, which are then identified as the group labelled "Euchelicerata". ) A 2019 analysis nests Xiphosura deeply within Arachnida. Pycnogonida (sea spiders) [REDACTED] Xiphosura (horseshoe crabs) [REDACTED] Arachnida [REDACTED] Discovering relationships within
2254-432: The terrestrial lifestyle of arachnids, such as internal respiratory surfaces in the form of tracheae , or modification of the book gill into a book lung , an internal series of vascular lamellae used for gas exchange with the air. While the tracheae are often individual systems of tubes, similar to those in insects, ricinuleids, pseudoscorpions, and some spiders possess sieve tracheae, in which several tubes arise in
2303-517: Was first used by Tamerlan Thorell in 1883. It means "tail rump", from Ancient Greek οὐροπύγιον ( ouropugion ), from οὐρά ( oura ) "tail" and πυγή ( puge ) "rump" referring to the whip-like flagellum on the end of the pygidium , a small plate made up of the last three segments of the abdominal exoskeleton . The classification and scientific name used for whip scorpions varies. Originally, Amblypygi (whip spiders), Uropygi and Schizomida (short-tailed whipscorpions) formed
2352-498: Was not included in the study), received strong support. Somewhat unexpectedly, there was support for a clade comprising Opiliones , Ricinulei and Solifugae , a combination not found in most other studies. In early 2019, a molecular phylogenetic analysis placed the horseshoe crabs, Xiphosura , as the sister group to Ricinulei. It also grouped pseudoscorpions with mites and ticks, which the authors considered may be due to long branch attraction . The addition of Scorpiones to produce
2401-493: Was the first to use a genus name solely for whip scorpions, namely Thelyphonus . Latreille later explained the name as meaning " qui tue ", meaning "who kills". One name for the order, Thelyphonida, is based on Latreille's genus name. It was first used, as the French Thélyphone , by Latreille in 1804, and later by Octavius Pickard-Cambridge in 1872 (with the spelling Thelyphonidea). The alternative name, Uropygi,
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